Animal Weapons

by Douglas J. Emlen

  Gorbachev, Mikhail

  grackles, great-tailed

  grasshoppers, foam

  Greeks, ancient

  Grenada

  grenades

  Grimaldi, David

  guerrilla warfare

  gun ports

  gunpowder

  habitat

  hares, snowshoe

  Hayden, Thomas

  health

  hedgehogs

  helmets

  Hiroshima and Nagasaki

  Hoekstra, Hopi

  Homer

  horns

  antelope

  beetle

  clades and

  duels and

  giraffes and

  ibex

  increases vs. decreases in

  Kyptoceras

  pig

  resources and

  sexual selection and

  Synthetoceras

  topi

  trilobite

  woolly rhinoceros

  horses, knights and

  horseshoe crabs

  humans. See also knights; soldiers; and specific battles; nations; and weapon types

  access to resources and

  armor and

  arms races between rival states and

  arms races ended with cheap weapons

  comparing animal weapons and

  costs of weapons and

  cultural vs. biological evolution and

  defensive weapons and

  deterrence and

  division of labor and

  duels and

  eggs vs. sperm size and

  male competition for females and

  Hunt, John

  hunter-gatherers

  hunting skills

  hyenas

  hyper-carnivore predators

  ibex

  improvised explosive devices (IEDs)

  Incan empire

  Incisitermes

  incisors

  India

  insects. See also specific types

  colonies, and body shape

  exoskeletons

  intercontinental ballistic missiles (ICBMs)

  Iran

  Iraq War

  Irish elk (Megaloceros giganteus)

  Iroquois

  Israel

  Italy

  jacanas

  Japan

  javelina

  jaws

  alligator gar

  ant

  caiman

  cat

  crocodile

  eel

  fish

  hyena

  piranha

  predator

  sabertooth

  speed vs. strength of

  wolf

  Jericho

  Judea

  K-324 submarine

  katydid

  leaf-mimicking

  spines

  Kaufman, Donald

  Kevlar

  kiwis

  knights

  knives

  Koga, Tsunenori

  Korean Air Lines Flight 007

  Korean War

  Kyptoceras

  Lachish

  La Milpa

  lances

  Lanchester, Frederick William

  Lanchester’s linear law

  Lanchesters’s square law

  land mines

  Latvia

  law of the lever

  leaf-footed bugs

  leaf mimicking

  Levinton, Jeff

  lions

  lizards

  lock-and-release mechanism

  longbows

  Lorch, Patrick

  Los Alamos Laboratories

  lures

  lynx

  Canada

  tooth size

  M1 rifle

  M4 rifle

  M16 rifle

  M1918 Browning Automatic Rifle (BAR)

  Mach II technology

  machine guns

  Maginot Line

  mammals. See also specific types

  predators

  mammoths

  Columbian

  woolly

  mandibles, stag beetle

  mantis shrimp

  manufactured structures

  Maori villages

  marine isopod crustaceans

  marsupials, sabertooth

  mastodons

  matted hair

  Mayan fortifications

  McCloy, USS (frigate)

  McElligott, Alan

  Me 262 fighter plane

  metabolism

  metal blades

  mice

  burrows

  field

  inland vs. beach

  microscopic weapons

  Middle Ages. See also knights

  MiG-15 fighter planes

  MiG-21 and MiG-23 fighter planes

  missiles

  Missouri, USS (battleship)

  mitochondria

  MKb 42 (H) rifle

  Moen, Ron

  Moffett, Mark

  molars

  Montserrat

  moose

  Gallic

  mountain lions

  Mullen, Lynne

  multiple independent reentry vehicles (MIRVs)

  muscle cells

  muskets

  mutations

  Napoleonic Wars

  NASA

  Nasutitermes soldiers

  NATO

  natural selection

  camouflage and

  carnivore teeth and

  fish jaws and

  sexual selection vs.

  stickleback spines and

  Nautilus, USS (submarine)

  Nicaragua

  Nile Valley

  nimravids

  North Korea

  nuclear weapons

  nutrition

  ogrefish

  Olmec empire

  Operation Enduring Freedom Camouflage Pattern (OCP)

  Oppenheimer, J. Robert

  osteoporosis

  owls

  P-51D Mustang fighter plane

  pair bonds, long-term

  Pakistan

  pangolins

  parasites

  parental care

  Pastor, John

  Patek, Sheila

  pathogens

  biological weapons and

  Patriot Advanced Capability antimissile system

  Pax Americana

  Pax Britannica

  Pax Romana

  peacocks

  peasant fighters

  pedipalps

  performance, balance between size and

  Pershing II missiles

  Petrov, Stanislav

  Phoenicians

  pigs

  pill bugs

  piranha

  points

  political arms races

  pollution

  Pomiankowski, Andrew

  Poole, Joyce

  porcupine fish, long-spine

  porcupines

  praying mantises

  predators

  ambush

  camouflage and

  fiddler crabs and

  hunting weapons and

  lake vs. marine habitats and

  stalking

  teeth of

  premolars

  pseudoscorpions

  Ptolemy V

  reproduction. See also sexual selection

  resources

  cost effectiveness and

  deterrence and

  discretionary pools within species and

  dung beetles and

  fiddler crabs and

  fixed

  localized, rare, and economically defensible

  political arms races and

  sneaky males and

  trade-offs and

  rhinoceros

  woolly

  Ribeyrolle 1918 rifle

  rifles

  RNA

  rocket technology

  Romans, ancient

>   ruffs

  Russia

  sabertooths

  salmon

  samurai

  satellite males

  Schluter, Dolph

  Schöning, Caspar

  scimitar-toothed cat

  scorpions

  scrambles

  seals, elephant

  Semna

  sexual selection

  arms race collapse and

  balance between costs and benefits and

  cicada-killer wasps and

  clades and

  competition and

  costs of weapons and

  differences within species and

  duels and

  economic defensibility and

  elephants and

  extreme

  female choice and

  harlequin beetles and

  horseshoe crabs and

  jacanas and

  knights and

  natural selection vs.

  New Guinean antlered fly and

  pseudoscorpions and

  sneaky males and

  stag beetles and

  stalk-eyed flies and

  weapon size enhanced by

  sheep, bighorn

  Sheridan tank

  shields

  ships

  aircraft carriers

  battleships

  bireme (“twos”)

  carrier strike groups

  Cold War

  container

  deterrence and

  destroyers

  dreadnoughts

  fire ships

  first-rate

  fives

  forty

  galleons

  ironclads

  Lanchester’s models and

  oared galleys

  penteconters

  Q-ships

  steam-powered

  trireme (“threes’)

  shrimp

  pistol

  siafu. See ants, African army

  siege warfare

  signals

  Simmons, Leigh

  sloths, giant

  Smilodon fatalis

  snakes

  sneaky tactics (cheating; sneak attacks)

  arms race collapse and

  bighorn sheep and

  costs vs. benefits and

  cuttlefish and

  dung beetles and

  evolution of

  humans and

  marine isopod crustaceans and

  ruffs and

  salmon and

  ships and

  sunfish

  soldier insects

  soldiers. See also knights; peasant fighters; and specific weapons

  artillery and

  body armor and

  camouflage

  Sopwith Tabloid biplane

  South Africa

  Soviet Union

  space race

  Spanish Armada

  spears

  Special Forces

  speed

  sperm

  spiderwebs

  spikes and spines

  Spratt, Norman

  spurs, jacana

  Sputnik

  squad automatic weapons (SAWs)

  squirters

  stalker fish

  star fort

  Stg 44 rifle

  sticklebacks

  three-spined

  Su-15 fighter planes

  Su-30 Flanker planes

  submarines

  suicide bombers

  Sumner, Francis Bertody

  sunfish

  Sun Tzu

  Swallow, John

  swine flu

  Synthetocreas

  Syracusans, ancient

  Taliban

  tanks

  teeth

  ambush predators and

  breakage

  dinosaur

  fish and

  mammalian predator

  omnivore

  trade-offs and

  Teleopsis

  dalmanni

  quinqueguttata

  whitei

  Tenochtitlan

  Terminal High Altitude Area Defense system

  termites

  fortifications

  Macrotermes subhyalinus

  queens

  soldiers

  terrorism

  testes

  theropod dinosaurs

  tigers

  Tigris-Euphrates rivers

  Tikal

  Titan II ICBMs

  titanotheres

  Titan Rain attacks

  toads, cane

  Tomkins, Joe

  topi

  torpedoes

  “Tortoise and the Hare, The” (Aesop)

  toxins

  trade-offs. See also costs vs. benefits; resources

  division of labor and

  portability and

  resource allocation and

  speed and

  teeth and

  trebuchets

  triceratops

  trilobites

  tuna

  tunnels

  side

  Turner, Ted

  turtles

  tusks

  Columbian mammoth

  dolphin

  elephant

  mastodon

  narwhal

  resources and

  ungulate

  wasp

  Tyrannosaurus rex

  Ukraine

  underground fortifications

  ungulate clades

  United States

  Cold War and

  cyberattacks on

  U.S. Army

  U.S. Navy

  Universal Camouflage Pattern (UCP)

  unmanned aerial vehicles (UAVs, drones)

  Uruk

  V-22 Osprey tilt-rotor aircraft

  vehicular arms races

  Vietnam War

  viperfish

  viruses

  Vulpavus

  walruses

  Warsaw Pact

  Washington, Lake

  wasps

  cicada-killer

  West African

  weapons of mass destruction

  weevils

  Weibel M/1932 rifle

  whales

  Wilkinson, Gerald

  wings

  beetle horns and

  insect soldiers and

  wolves

  World War I

  World War II

  Wright, Orville and Wilbur

  Younger Dryas

  zebra

  Zeh, David

  Zeh, Jeanne

  Zeleny Island airspace

  zero-day attacks

  The earliest human paintings—some more than 30,000 years old—portray animals with extreme weapons. Wooly rhinoceros from Chauvet Cave (Above) and Irish elk from Lascaux Cave (Below).

  Elk pay a steep price for their weapons. Bulls double their daily energetic needs and leach essential minerals away from other bones in order to grow antlers.

  Many animals, such as this katydid, rely on camouflage to avoid being eaten by predators. Leaf-mimic katydids even rock back and forth when they walk, resembling a leaf fluttering in the wind.

  Special Forces snipers also rely on camouflage, choosing from among a selection of specialized suits the one most likely to match the particular background of each mission. Exquisite background matching is not cost effective for most other situations, and the majority of soldiers wear uniforms that provide only moderate camouflage—a universal color pattern designed to work reasonably well against a range of different backgrounds.

  Animals employ diverse defensive weapons, including spines and armor.

  Cats are agile predators whose weapons are moderate in size, reflecting a balance between selection for killing large prey and selection for agility and speed. Above: Mountain lions behind the author’s house. Below: Canada lynx.

  Weapons in ambush predators experience very different selection from weapons in other predators. Inst
ead of running, swimming, or flying fast to catch prey, ambush predators snatch them with a quick strike from claspers. As a result, many have large weapons. Above: Mantisfly. Below: Peacock mantis shrimp.

  The most extreme animal weapons are wielded by males, and used in battles for access to females. Above: Rhinoceros beetle horns. Below: Harlequin beetle forelegs.

  Diversity in dung beetle horns. All of these species are “tunnelers,” who use their weapons in fights for burrows containing females.

  When males fight in unrestricted places, such as in the air, or in chaotic scrambles, agility takes precedence over big weapons. Above: Cicada killer wasps. Below: Horseshoe crabs.

  Males in many animals, such as these flies, fight over restricted, economically defensible resources visited by females. In these situations, big weapons are often beneficial. Above: Stalk-eyed flies defend hanging rootlets on which females roost. Below: Antlered flies defend egg-laying holes in the bark of fallen trees.

  Duels are an essential ingredient of an arms race. When rival males face off one-on-one, selection often favors the male with the bigger weapons, driving evolution of extreme weapon sizes. Above: Stag beetle males. Below: White-tailed deer.

  Vehicles also often fight in one-on-one duels and, as in animals, this can trigger an arms race. Above: The addition of a battering ram precipitated explosive growth in ship size in oared galleys of the ancient Mediterranean, settling in the end on the “five.” Below: Side-mounted cannons had the same effect on sailing galleons.

  Fallow deer and moose pay exorbitantly for their weapons, and for the stamina and energy needed to win fights during the rut, suffering gashes, infections, and depleted energy reserves. Above: Three-quarters of fallow deer bucks die without ever succeeding in defending territories, and 90 percent fail to mate even once in their lifetime. Below: Bull moose double their daily energetic demands while growing antlers, and a third die from injuries sustained during combat.

  Battling for access to females is dangerous, resulting frequently in serious wounds and, occasionally, death. Above: Fighting male oryx. Below: Male antelope died after their horns locked together in combat.

  Relative to their body size, male fiddler crabs wield the largest weapons of any animal.

  Because fighting is dangerous, males assess each other beforehand, comparing weapons and pushing. Claws function as deterrents, and fights only escalate when males are evenly matched.

  Ships function as deterrents, too. Above: Capital “ships of the line” were the most expensive and powerful weapons of their day. Out of reach to all but the richest nations, these ships projected power to the farthest reaches of the globe. Below: Nimitz-class aircraft carriers function in this same capacity today.

  Searching for sneaky males. Above: Male dung beetles fight inside a tunnel; the female is beneath them to the right. Below: Observing behavior inside tunnels is possible using “ant farms.” Small, hornless males sometimes sneak into guarded tunnels below ground, digging side tunnels that intercept the main tunnels beneath guarding males.