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ANTS
ANTS
A VISUAL GUIDE
HE ATHER C A MPBELL A ND BENJA MIN BL A NCH A R D

PR I N C ETON UNIV ER S ITY PR ES S


PR I NC ETON AND OX F OR D
Published in 2023 by Princeton University Press
41 William Street, Princeton, New Jersey 08540
99 Banbury Road, Oxford OX2 6JX
press.princeton.edu

Copyright © 2023 by Quarto Publishing plc

All rights reserved. No part of this book may be reproduced or transmitted in any form or by
any means, electronic or mechanical, including photocopying, recording or by any information
storage-and-retrieval system, without written permission from the copyright holder.

Library of Congress Control Number 2022940432


British Library Cataloging-in-Publication Data is available

ISBN: 978-0-691-22852-5

This book was conceived, designed, and produced by


The Bright Press, an imprint of the Quarto Group,
The Old Brewery, 6 Blundell Street,
London N7 9BH, United Kingdom.
T (0)20 7700 6700
www.quarto.com

The Bright Press


publisher: James Evans
editorial director: Isheeta Mustafi
art director: James Lawrence
managing editor: Jacqui Sayers
project editors: Joanna Bentley, Anna Southgate
design: Kevin Knight
picture research: Jane Smith

Cover images: (front) Adisak Mitrprayoon/istock;


(spine) Simon Dannhauer/Shutterstock
Cover design by: Kevin Knight

Typeset in: Chaparral Pro

Printed and bound in Singapore

10 9 8 7 6 5 4 3 2 1
CONTENTS

Introducing the Ants 6

chapter 1 chapter 3 chapter 5


W H AT I S A N A N T ? 14 L I FE H I S TO RY, E CO LO GY 150
Ant Anatomy 16 R E P RO D U C T I O N, The Hallmark of Ant Ecology 152
When Form Meets Function 18 A N D D E V E LO PM E N T 80 Finders, Keepers . . . Or Not? 156
Attack and Defense 20 Life Cycle 82 Symbiotic Living 160
Locomotion 22 Love is in the Air 85 No Harm, No Foul 164
Sensory Structures 24 Colony Foundation Prowling Hunters
Internal Anatomy 26 —Variations on a Theme 88 and the Enemy Within 167
Reproduction 28 Sex Determination and Eusociality 92 Ants in the Anthropocene 170
Ant Profiles: Castes in Ants 94 Ant Profiles:
Odontomachus 30 Worker Exploitation 100 Azteca 172
Gigantiops 32 Ant Profiles: Rhytidoponera 174
Polyrhachis 34 Mystrium 102 Anoplolepis 176
Carebara 36 Stigmatomma 104 Camponotus 178
Melissotarsus 38 Eciton 106 Atta 180
Harpegnathos 40 Formica 108 Neoponera 182
Tatuidris 42 Oecophylla 110 Tetraponera 184
Messor 112
chapter 2 chapter 6
Dinoponera 114
E VO L U T I O N ANTS AND PEOPLE 186
A N D D I V E RS I T Y 44 chapter 4 Farming Friend or Foe? 188
The Ant Tree of Life 46 B E H AV I O R 116 Food and Pharmaceuticals 190
The Big Five 48 Trailblazing Work 118 Cultural Icons 194
Rise to Dominance 52 Colony Chatter 122 Artistic Inspiration 196
Deep Time Capsules 54 Spit Takes 124 School of Ants 198
Innovation and Trade-offs 56 Seeing is Believing 126 Change Underfoot 200
A Cosmopolitan Group 59 Safe and Secure 128 Ant Profiles:
Island Living 62 Builders and Architects 131 Linepithema 202
Ant Profiles: Memory in Time 134 Dorylus 204
Nothomyrmecia 66 Ant Profiles: Lasius 206
Crematogaster 68 Cataglyphis 136 Melophorus 208
Pheidole 70 Colobopsis 138 Myrmecia 210
Strumigenys 72 Euprenolepis 140 Tetramorium 212
Haidomyrmecinae 74 Aphaenogaster 142 Paraponera 214
Martialis 76 Cephalotes 144
Discothyrea 78 Megaponera 146 Glossary 216
Plectroctena 148 Further Reading 219
Index 221
Picture Credits 223
Acknowledgments 224
INTRODUCING THE ANTS

THE HUMBLE ANT IS HARDWORKING, UBIQUITOUS, and specialized plant structures, to their role tending sap-sucking
instantly recognizable to people across the globe. From city bugs. They are voracious predators renowned for their
sidewalk to rainforest canopy, we cross paths with ants, yet we strength and ability to take down prey much larger than
rarely consider what has made them so successful that they themselves using chemical weapons alongside impressive
occupy virtually every nook and cranny of land on Earth. Ants jaws and even more impressive stings. Ants host a range
can make up one-third of insect biomass, an amount many of visitors in their nests, from other invertebrates to fungi
times greater than all terrestrial vertebrates. A single colony that they cultivate for food. Some creatures, such as spiders,
of Leafcutter Ants has more individuals than the entire mimic them hoping to gain a benefit from appearing to be
population of New York City (10 million ants vs. 8.6 million an aggressive or unpalatable ant.
people). But profusion alone does not define their importance;
rather, it is their essential roles in ecosystems and their vast As a group, the Formicidae—better known as ants—
host of interactions with other organisms. Take the Army Ant are a family that sits within the larger insect order of
Eciton burchellii as an example; this single species is associated Hymenoptera, which includes bees, sawflies, and wasps.
with at least 557 other species, including over 200 birds. We know of around 14,000 ant species, but estimates add
potentially another 10,000 species not yet discovered or
Their relationships with other species cover a vast range of described by taxonomists. Belonging to a single family of
interactions—from tightly coevolved mutualisms nesting in insects, ants are subdivided into 16 living subfamilies with

L E F T: A widespread ant–insect
mutualism is the tending of
phloem-feeding hemipterans,
which produce sugary
honeydew. The ants feed on this
and in return provide protection
against predator and parasitoid
attack for the bugs.

OPPOSITE TOP: The


possession of elaborate spines
is part of an evolutionary
trade-off against other ant
defensive traits, such as having
a functional sting.

OPPOSITE BOTTOM: Worker


ants, such as these Leptogenys,
will move eggs, larvae, and
pupae within a nest and to
new nest sites.

6 INTRODUCING THE ANTS


mate with a queen. In true ant style, this insect has developed
many variations on this basic life history. Some species have
reproductive workers instead of a queen caste, while others
live parasitically inside the nest of host species.

Their behaviors are perhaps the thing that amazes us


most about ants: They have a remarkable ability to cooperate
and communicate to defend their nest from intruders or
to collectively hunt and bring back food to their nest. The
reproductive capacity of a queen ant sees her producing
an additional 6 subfamilies that contain only extinct species. over 250 million eggs in a lifetime. And a queen is capable
Diversity is highly variable across subfamily groups: some of invading the nest of another species, killing the original
contain a single species while the hugely successful queen, and then manipulating the remaining workers of an
Myrmicinae subfamily has over 7,000 species. entirely different species to feed and raise her young for her.
We even know of ants who rescue their injured nestmates
While it is easy enough to point out an ant when we see and act as ambulances, carrying the injured back to the nest
one, it is harder to pinpoint what exactly makes it an ant. to rest and recuperate. All of these amazing facets of ant
Ants are characterized by modified abdominal waist behavior, alongside their interactions, anatomy, and life
segments, elbowed antennae, and a metapleural gland. More history, are covered in this book.
recognizable is the lack of winged individuals in the worker
caste, which sets them apart from many of their nearest bee
and wasp relatives. By eye they generally appear to be dark
six-legged specks that are fairly hard to tell apart while busily
scurrying around. Under a microscope they are revealed to
have huge variation in their appearance, with features such
as scythe-like snapping jaws, ostentatious spikes, bizarrely
shaped hairs, or large glitter ball eyes.

We find ourselves fascinated by ant societies—female-led


colonies where members work cooperatively to find food and
raise young. Colonies may contain a few individuals up to a
million, depending on the species. Colonies have one or more
queens, with daughter workers who forage for food and take
care of the young within the nest. Effectively this means
that all workers are closely related as sisters. Queens can be
long-lived (up to 30 years in one species), in contrast to males,
who are generally short-lived, existing only long enough to

INTRODUCING THE ANTS 7


A N T C L A SS I FI C AT I O N C H A RT

Phylum: Arthropoda | Class: Insecta | Order: Hymenoptera | Family: Formicidae

Subfamilies Diversity Notes

The “Big Five”

Dolichoderinae 28 genera, 714 species (136 species) Single petiole and slit-like orifice used for chemical defense.
Most speciose genera are Dolichoderus, Technomyrmex,
Azteca, and Iridomyrmex.
Dorylinae 27 genera, 749 species (8 species) Robust sting. Most speciose genera are Aenictus,
Neivamyrmex, and Lioponera.
Formicinae 52 genera, 3,244 species (171 species) This subfamily contains 23 percent of the world’s ant species.
Single petiole and acidopore. Over a thousand species
are Camponotus.
Myrmicinae 147 genera, 7,075 species (187 species) Over half of the world’s species are in this subfamily.
Petiole and postpetiole nodes. Pheidole, Strumigenys, and
Tetramorium are most speciose genera.
Ponerinae 50 genera, 1,267 species (87 species) Single petiole segment and usually a robust sting.
Most diverse genera are Leptogenys, Hypoponera,
and Anochetus.

Smaller Subfamilies

Agroecomyrmecinae 2 genera, 2 species (3 species) 2 living species, Ankylomyrma coronacantha and Tatuidris
tatusia
Amblyoponinae 9 genera, 143 species (7 species) Highest diversity in Fulakora, Stigmatomma, and
Prionopelta
Aneuretinae 1 genus, 1 species (12 species) Single living species, Aneuretus simoni, from Sri Lanka
Apomyrminae 1 genus, 1 species 1 species, Apomyrma stygia, from West Africa
Ectatomminae 12 genera, 302 species (14 species) Most diverse genera are Gnamptogenys and Rhytidoponera
Leptanillinae 7 genera, 70 species Most diverse genus is Leptanilla
Martialinae 1 genus, 1 species 1 species, Martialis heureka, from Brazil
Myrmeciinae 2 genera, 94 species (22 species) All living species are in Myrmecia (93 spp.) except 1 species
in Nothomyrmecia
Paraponerinae 1 genus, 1 species (1 species) 1 living species, Paraponera clavata, and 1 fossil species,
Paraponera dieteri
Proceratiinae 3 genera, 165 species (12 species) 3 genera are Proceratium, Discothyrea, and
Probolomyrmex
Pseudomyrmecinae 3 genera, 235 species (20 species) 3 genera are Pseudomyrmex, Tetraponera, and the single-
species genus Myrcidris

8 INTRODUCING THE ANTS


Subfamilies Diversity Notes

Known only from fossils

Armaniinae 2 genera, 12 species Genera are Armania and Pseudarmania


Brownimeciinae 1 genus, 1 species Monotypic genus with 1 species, Brownimecia clavata
Formiciinae 2 genera, 6 species Genera are Formicium and Titanomyrma
Haidomyrmecinae 10 genera, 16 species All genera have 1 species except for Ceratomyrmex (2 spp.),
Haidomyrmex (4 spp.), and Linguamyrmex (3 spp.)
Sphecomyrminae 4 genera, 12 species Gerontoformica is the most diverse genus (9 species)
Zigrasimeciinae 3 genera, 6 species Genera are Boltonimecia (1 sp.), Protozigrasimecia (1 sp.), and
Zigrasimecia (4 spp.)

Figures listed give only the extant diversity within a subfamily. Figures in parentheses give the additional diversity for extinct species.
The diversity within subfamilies known only from fossils are all extinct genera and species.

W H O STU D I ES A NTS?
Scientists who specialize in conducting research on ant biology
are known as myrmecologists, with the field of ant biology
itself referred to as myrmecology (from the Greek myrmex for
ants). As well as natural scientists who study many aspects of
the biology of ants, computer scientists and engineers also
study these insects. Research hubs exist in the USA, Brazil,
Germany, Australia, France, Japan, China, and the UK. As
with many subdisciplines of entomology, tropical regions are
generally less well characterized in terms of their ant fauna
even though they have the greatest species diversity. This is
changing, but requires investment in ant research for countries
in the Global South. Traditionally, European and American
men have been credited as pioneers in myrmecology with
contributions from women and scientists from other countries
often being overlooked. Currently, women represent around
one third of authors publishing research on ants, and data is
not available on the proportion of scientists from other
marginalized groups. While participation is increasing, there
is still a long way to go to create a truly diverse and equitable ABOVE: Within a nest, adult worker ants are found alongside
community of myrmecologists. a queen as well as juvenile stages of the colony known as the
brood (eggs, larvae, and pupae).

INTRODUCING THE ANTS 9


A NT EN VI RO N M ENTS Oceania includes Micronesia, Fijian islands, Hawaiian
Ants are found in all parts of the globe except the coldest islands, and Polynesia. The final zone, the Antarctic, is not
polar regions and some islands. Throughout the book we occupied by ants. Across all regions most terrestrial habitats
use eight broad biogeographic regions to describe the are occupied by ants. These can include natural and semi-
distributions of different species. The Neotropical realm natural forest, woodland, grassland, scrub, wetland, and
includes Central and South America as well as the Caribbean, desert areas. Ants frequently occupy human-modified areas,
with the Nearctic covering most of North America. The vast such as agricultural land used for crops, livestock pasture
Palearctic region comprises Europe, Asia from north of the and forest plantations, or urban spaces.
Himalayas, and Africa northwards of the Sahara. The
Holarctic is sometimes used to collectively refer to a Northern
Hemisphere distribution of ants that combines the Nearctic BE LOW: Ants are adapted to OPPOSITE: The seven
and Palearctic regions. The Afrotropical realm includes Africa nesting in many microhabitats biogeographic realms that
south of the Sahara, parts of the Arabian Peninsula, and the above and below ground. They are used throughout this
island of Madagascar. The Indomalayan region is made up are especially influenced by the book when describing the
of India and Southeast Asia, while Australasia includes availability and structure of live distribution patterns of ants
Australia, New Guinea, and the eastern part of Indonesia. and dead vegetation. (Antarctica excluded).

A N T N ES TI N G SITES

Woven from leaves


Ant garden from
plant material and
epiphytic plants

Excavated
Mound live wood
building
Under logs or
inside rotting/
In soil hollow logs
under stones

10 INTRODUCING THE ANTS


The majority of ants occupy a nest or series of nests where Arboreal ants nest on or in trees, exploiting a host of strategies.
the colony lives, providing safe and stable conditions for Some ants can tunnel into live wood, creating their own holes
the queen and brood, as well as a central point from which to nest in or excavating a network of galleries under bark.
workers forage for food. Other ants have a nomadic lifestyle, Others are able to weave together leaves to construct a nest
foraging and nesting on the move, and sometimes constructing or create ant gardens, a creation that combines recycled plant
bivouac shelters weaved from their own bodies. material with epiphytic plant seeds sown in for structural
integrity. Plants may provide domatia, specialized modified
Ants broadly fall into two nesting categories: ground or structures like swollen thorns or hollow branch segments
arboreal. Ground-nesting ants have a host of options. They that are specifically grown for ant occupation.
may be subterranean, excavating their own nests deep
within the soil; live in leaf-litter; or nest opportunistically Ants often show a specialization to a particular foraging or
under rocks and logs. Ground-nesting ants make use of plant nesting stratum (subterranean, leaf litter, canopy) but are not
material even when they do not directly occupy trees. They strictly limited to foraging in the same microhabitat that they
will nest in cavities such as hollow dead branches and twigs, nest in. Many arboreal nesting ants will forage on the ground,
or inside nuts and seeds on the forest floor, and especially while ground-nesting ants move on to trees to hunt or find
inside rotting logs. plant-based resources like nectar or honeydew from insects.

M A P O F BI O G EO G R A PH I C RE A L MS

Palearctic
Nearctic

Indomalayan
Afrotropical
Division of the
biogeographic realms Oceanian
approximately follows
the red dashed lines. Neotropical
Australasian

Areas shaded green are


known to contain high species
diversity that is generally
concentrated in tropical forests.

INTRODUCING THE ANTS 11


A BO UT TH IS BO O K Ecology undertakes a review of the main topics in ant
This book contains six chapters covering different aspects ecology—an area with a substantial amount of scientific
of the biology and ecology of ants. While many topics are research. We begin by examining direct and indirect
intertwined, this organization allows us to explore major competitive ant interactions and how these relate to ant
themes in the natural history of ants. Accompanying the coexistence and niche partitioning. We then look at a range
topic chapters are profiles of ant genera that highlight the of ant interactions with other organisms such as mutualisms,
unique stories of the featured groups. commensalisms, and antagonistic interactions with predators
and parasites. Finally, we briefly cover the impact of human-
TO PI C C H A P TERS induced habitat loss and routes of ant species conservation.
What is an Ant? offers an overview of the physical features
of ants, including their basic body plan. We examine details Ants and People shows how ants touch human lives in diverse
of external anatomy, including morphology used in attack ways, for example through their role in agriculture, their use
and defense, as well as structures used for locomotion and as food and pharmaceuticals, and the cultural value of ants in
sensing the environment. We also look at the internal art and storytelling. We end by describing how the public has
systems of ants and outline their reproductive anatomy. been involved in ant research through citizen science and the
global impacts of invasive ant species.
Evolution and Diversity summarizes ant evolution and
examines in turn each of the “Big Five” most diverse ant A NT PRO FI L ES
subfamilies. We look at theories explaining ant dominance, The ant profiles are an opportunity to delve deeper into the
and the innovations and trade-offs that have contributed to biology of a particular group. These are typically presented
their success. The fossil record for ants is discussed, and then at the level of genus, with the exception of the fossil group
we highlight the global distribution patterns of ant diversity, of Hell Ants, which are a subfamily. Common names are
especially ants found on islands. also given where they exist, although many ants do not have
them, and other genera share a common name. Each profile
Life History, Reproduction, and Development gives an insight gives a brief overview of the biology and ecology before
into the ant life cycle, including how and when mating occurs. focusing on interesting features of the taxon. Due to ants’
Colony foundation occurs in many ways for ants, and these small size and cryptic lifestyles, many aspects of their biology
are discussed followed by the process of sex determination. are not well understood, but the detail in these profiles gives
Eusociality and the different castes of ants are outlined, with some glimpse into what we currently know about them.
a discussion on exploitation of workers. Information on distributions, habitats occupied, and feeding
and nesting preferences are given to the best of current
Behavior begins by introducing how ants navigate and scientific knowledge.
forage and goes on to outline the varying methods of ant
communication. There then follows a description of the ant
“social stomach” and trophallaxis, followed by a review of OPPOSITE: Among the topics discussed are (clockwise from top left)
other diverse behaviors, including the use of vision, external ant anatomy; morphological traits; communication; impact on
aggression, defense, nest construction, and memory. the environment; interactions in the natural world; and colony migration.

12 INTRODUCING THE ANTS


INTRODUCING THE ANTS 13
1 W2
H A T I3
S A N4A N T5
? 6
A N T A N ATOMY

ANTS CAN BE CHARACTERIZED BY a combination of three M ESOSOM A


unique physical features: a distinct one- or two-segmented The size of the mesosoma relative to the rest of the body
waist, elbowed antennae, and a metapleural gland. They is a good indicator of whether an ant will have wings and
follow a basic insect body plan. This includes six legs, two fly at any stage of its life cycle. In the case of males and
pairs of wings (which are absent in workers), and division into reproductive females, queens, the thorax is enlarged to allow
three main body sections. As invertebrates they have no spine space for the flight muscles. Where present, ants possess
or internal skeleton, instead relying on a hard, waterproof two pairs of membranous wings. Wingless ants may be
exoskeleton made of chitin to give their body rigidity and queens who have lost their wings after use—who are dealate.
protection. The exoskeleton is punctuated with spiracles, Alternatively, they may be female workers who never possess
small breathing holes that allow the uptake of oxygen into them to begin with. They are generally easy to tell apart as
the body. Beyond this superficially simple anatomical dealate queens are larger than workers and possess wing
outline, ants exhibit a wondrous range of physical adaptations. scars where wings have been bitten or rubbed off.
It is this specialization of form and function that has allowed
ants to flourish in virtually every terrestrial habitat on Earth. GASTER
When it comes to organs, the ant brain and associated
BO DY PL A N nervous system, along with the heart, run longitudinally
Insects have three body regions: the head, thorax, and through the mesosoma and gaster. Within the gaster, ants
abdomen, whereas ants have differently named sections: the house their digestive and reproductive organs. They have two
head, mesosoma, and gaster. The mesosoma is the middle stomachs, one for their own digestion and the other a social
body section, formed as the first abdominal segment (the stomach for sharing food in the colony. At the tip of the
propodeum) is fused to the thorax. The next one or two gaster, external evidence of defensive structures can be seen.
abdominal segments are much reduced in size to become Depending on the ant subfamily, they may possess a sting, a
small nodes. They form a distinctive waist known as the formic acid–spraying funnel of hairs known as an acidopore,
petiole and (if present) postpetiole, which is a defining or a simple slit. A unique identifying character of ants is the
characteristic of ants. These waist segments are useful to presence of the metapleural gland, although it has been lost
prevent misidentification where ants might be confused with in some subfamilies. This slit or round-shaped opening in
small wasps or the erroneously named Velvet Ants, which are the exoskeleton is a secretory gland found at the base of the
not in fact ants at all. The first step in identifying an ant to propodeum. Its full significance is not understood. It could
subfamily is to count the number of waist segments. be for odor recognition or territorial marking but is most
Subfamilies including the Dolichoderinae, Formicinae, and likely used in sanitation and chemical defense.
Ponerinae have one waist segment, while the Myrmicinae
and Pseudomyrmecinae have two waist segments. Following All the structures mentioned in this overview are elaborated
the petiolar nodes is the gaster, composed of the remaining on in the sections that follow. The different subfamilies are
abdominal segments. treated in more detail in Chapter 2.

16 WHAT IS AN ANT?
A N T A N ATOMY

Petiole
Head Mesosoma Gaster

Mesonotum Propodeum
Pronotum

Spiracle
Metapleural gland

Clypeus
Acidopore
Mandible

ABOVE: As shown here a single waist segment, the petiole, is


typical of subfamilies including the Dolichoderinae, Formicinae,
and Ponerinae. Another notable feature includes the presence
of a metapleural gland, a unique structure found only in ants.

petiole
Petiole

Post-
Head Mesosoma Gaster

Propodeal spine
Antennal scrobe

Antennae

Scape

Funiculus Sting

Subpetiolar process
Compound eye
Leg

ABOVE: The presence of antennal scrobes allows an ant to retract


its antennae to be flush with its face. The two-segmented waist,
comprised of a petiole and postpetiole, is indicative of ants belonging
to subfamilies such as the Myrmicinae and Pseudomyrmecinae.

ANT ANATOMY 17
WHEN FORM MEETS FUNCTION

MANY ASPECTS OF AN ANT’S appearance have likely Australia is home to ants in an even wider array of colors:
evolved to meet a specific lifestyle requirement, although the Rhytidoponera metallica is more commonly known as the
extent to which this is true has not been fully explored for all Green-Headed Ant for its green-blue metallic appearance,
aspects of anatomy. Adaptations could be due to environment, while ants in the genus Iridomyrmex, meaning Rainbow
available food, or predators. Long legs and large eyes are Ant, often possess blue, green, or purple iridescence. The
commonly seen in ground-foraging ants that need to move most dazzling of all ants are those covered in a fine layer of
quickly to avoid predators in open ground or be the first to golden hairs, such as the spectacular Golden Carpenter Ant,
acquire a food resource. In contrast, ants that forage and nest Camponotus sericeiventris, found in forests across Central
in leaf litter have shorter legs and antennae, alongside small and South America.
eyes. This makes sense in the dark environment of leaf litter
where moving through small spaces is easier with a compact The thick cuticle that forms the ant exoskeleton can be highly
body plan. Based on the unique combination of body size patterned and covered in hairs. In fact, the terminology alone
measurements, scientists can predict where an ant nests to describe “sculpturation”—the depressions and ridges of the
and forages or even what kind of food it eats. Predators have cuticle—requires a 31-page guide. Ants range from having
longer, flatter mandibles, while omnivores—those eating a unsculptured, smooth, or glossy surfaces free of marks, to
diverse range of foods—have shorter, curved mandibles. superficial wrinkles and bumps, through to deep corrugations
and dense punctuations. The surface landscape of an ant sees
Size and shape alone are not the only ways in which ants sculpturing combined with “pilosity,” the density of rigid
vary in appearance. After seeing them under a microscope, hairs (setae) on the body, and “pubescence,” the shorter fine
the idea of ants as uniform specks of red or black quickly hairs elsewhere. Hairs on ants come in many different shapes,
evaporates. While they may not compare with the gaudy including clavate (club-shaped), ovate (egg-shaped), reniform
splendor of multicolored butterfly wings, they do exhibit a (kidney-shaped), and spatulate (spatula-shaped). Specialized
range of hues in blacks, browns, reds, oranges, and yellows. hairs may help defend ants against predators, while certain

BE LOW: Known commonly as Meat Ants, BE LOW: Calyptomyrmex piripilis is


Iridomyrmex purpureus have an orange-to-red found across the Afrotropics from
head and mesosoma with strong iridescence Sudan to South Africa. It has small
visible on their gaster. eyes and characteristic scale-shaped
hairs of unknown function.

18 WHAT IS AN ANT?
patterns of sculpturing can create a stronger cuticle, another
line of defense. Being hairy and having a thicker, more
sculptured cuticle can also help avoid desiccation through
water loss. It is even likely that these properties assist in
thermoregulation, as hair thickness and reflectivity may
help control body temperature and provide UV resistance.

L E F T: The widespread Australian genus


Rhytidoponera exhibits a range of metallic
colors from green and blue through purple.

B E L O W L E F T: Arboreal BE LOW: Found in Neotropical


Cataulacus ants have thick, highly rainforests, Golden Carpenter
sculptured exoskeletons, often Ants are covered in a layer
with protective teeth or spines on of long, dense golden (or
their thorax and propodeum. sometimes silver) hair.

WHEN FORM MEETS FUNCTION 19


AT TAC K A N D D E FE NS E

ANTS ARE NOTORIOUS TO GARDENERS and outdoors and curvature from small bumps to elaborate branched
folk alike for their ability to bite or sting at an unexpected structures. Considering the impressive spinescence of ants,
moment. They have an extensive arsenal of physical and the function is not studied in many species. In contrast to
chemical defenses at their disposal. Ants use hairs, spines, other ornate structures, such as the horns of Stag Beetles, ant
and teeth for attack and defense, yet their most lethal spines are not a product of sexual selection in males but are
weapon is located at the pointy end of the body. The most strongly displayed in the female workers. Unlike close
subfamily Formicinae have an acidopore, a conical structure relatives such as bees and wasps, wingless ant workers cannot
surrounded by hairs that squirts formic acid at any would-be fly away. It is thought that ant spines make it harder for
attackers. In contrast, the subfamily Dolichoderinae have vertebrate predators to capture and eat ants as well as for
a slit-like opening with no hairs or sting. Other major other invertebrates to attack them. Spines have evolved
subfamilies possess a venom-delivering sting for subduing
prey or defending themselves. An internal poison gland
BE LOW: Acanthognathus
produces either formic acid (in the Formicinae) or venom. B E L O W L E F T: This species ocellatus is a Neotropical
of Polyrhachis has three Trap-Jaw Ant that uses its rapidly
Ants have dorsal spines, protrusions of the cuticle on the separate pairs of defensive closing mandibles to catch
upper surface of the body. They usually extend from the spines on their pronotum, springtails and other small
propodeum, pronotum, or petiole. Spines range in length propodeum, and petiole. leaf-litter arthropods.

20 WHAT IS AN ANT?
repeatedly in many different and unrelated species, being for Daceton and Strumigenys, and in Ponerinae for
present in at least 79 of 346 genera, most commonly in the Anochetus and Odontomachus.
megadiverse subfamily Myrmicinae.
Some species have different worker castes, with one specifically
Jaws, or mandibles, are highly modified for feeding and in assigned a role in defense and usually referred to as soldier
predatory species especially for hunting. As well as mandibles ants. These are often larger than other workers in the colony.
being used in direct attacks, they can be used to hold a prey Another physical defense strategy is phragmosis. Phragmotic
item while a sting is deployed or to pin the prey in place while species have truncated body structures that are used to block
other workers dissect the unlucky individual. Trap-Jaw Ants the entrance to their nests. This is most commonly seen in the
have specialized mandibles that are held in an open position head of ants from genera such as Colobopsis, Cephalotes, and
and have trigger hairs that, when tripped, cause the jaws to Pheidole. Cephalotes, better known as Turtle Ants, have
shut with extreme force. While these jaws are used mainly in shield-like heads that can be used to plug the entrance to their
hunting, they can also be co-opted as a defense mechanism nest without exposing the eyes or antennae. In Cephalotes
to escape predators. Examples of Trap-Jaw Ants have varians, the head perfectly fits the nest entrance and the guard
evolved multiple times in different subfamilies, including in ant can allow other workers to enter and exit the nest while
Myrmoteras within Formicinae, in the Myrmicinae subfamily ensuring that any potential intruders are denied access.

BE LOW: Major Colobopsis A N T D EFENSE


etiolata worker ants have an B E L O W R I G H T: Defensive
abruptly truncated phragmotic structures include a sting (top left),
head. They use their head to an acidopore (top right), and a
block the entrance hole of nests slit-like opening used to smear
in dead branches or oak galls. chemicals (bottom).

Sting Acidopore

Slit-like opening

AT TACK AND DEFENSE 21


LOCOMOTION

A RELATIVELY SMALL SUBSET of all ants use flight to get MUSC L ES


around, with wings mainly present in males and queens. Queens have a morphology more typically seen of flying
Sometimes reproductive ants are wingless or show a insects, with an enlarged thorax to carry the musculature
reduction in wings (with short and long wings present in required for wings. In most flying insects the first thoracic
different morphs of the same species), but generally they have segment is small while the second and third are larger as each
two pairs of soft membranous wings. The requirement for supports a pair of wings. The thorax arrangement for queens
wings is because reproductives move much greater distances depends on their colony-founding strategy and therefore how
away from their birth nest in order to mate and in the case of they make use of their wings. Non-claustral species are those
queens to found a new colony. Workers move on foot, either where the queen will actively hunt to feed the first generation
to and from a central nest or with a colony in nomadic of workers. Queens perform a high amount of foraging and
species. They usually move over the ground or vegetation, their first thoracic segment is much closer in size to workers’,
often through confined spaces, none of which would be with their wing and neck muscles more balanced in size. In
feasible with two pairs of membranous wings. This means contrast, claustral queens do not forage or go through a
that the morphology of ants differs wildly depending on worker-like phase. They metabolize their own wing muscles
the caste of the individual being considered. to provide the energy needed for production of the first

T H O R ACI C MUSCL E A RR A N G E M EN TS

Dorsal cervical muscles

Wing muscles

Notopleural muscles

Winged queen Wingless worker

22 WHAT IS AN ANT?
generation of workers and therefore have a reduced neck L EGS
segment while the wing muscles are enlarged. In contrast Ants have three pairs of legs that vary in size and shape to
to the queens, worker ants show an enlarged first segment reflect the environment they occupy as well as their feeding
of the thorax and the muscles associated with it. This gives habits. These legs are named according to where they attach
greater strength and mobility to the neck, which is in control to the body. They are pro-, meso-, and meta-, referring to the
of head movements. first, second, and third pairs of legs. The basal segment where
the leg anchors to the body is the coxa. From there is found
the trochanter, a relatively small segment, followed by the
longer femur and tibia—equivalent to a human thigh and
shin. The final section is the tarsus, which has five
BE LOW: Ant legs can be highly
OPPOSITE: The structure of modified depending on how and
subsegments and terminates in a pair of claws—sometimes
thoracic muscles is very different where a species lives and forages, with a membranous lobe, the arolium, between the claws.
for winged queens (left) who have but they all follow the same The tibia sometimes possesses spurs that may be modified on
highly developed muscles for flight, broad plan for each leg of coxa, the forelegs as a specialized device for cleaning the antennae.
unlike wingless workers (right). trochanter, femur, tibia, and tarsus.

L EG S TRU C T U RES

Coxa Trochanter

Femur

Tibia

Tarsus

Tarsal claw Tibial spur

LOCOMOTION 23
SENSORY STRUCTURES

E YES chemicals. They can perceive UV light and have color vision,
Ants have two compound eyes composed of many facets although this is likely limited to only a few colors. Ants also
known as ommatidia. Eyes have been lost completely or process other visual information about motion, patterns,
reduced in many subterranean species, contrasting with a directions (position of the sun, polarization pattern of the
number of predatory species that have extremely large eyes skylight), and landmarks.
for hunting. The more ommatidia, the better the spatial
resolution of the ant’s vision. In ant species with large eyes, A NTEN N A E
ommatidia numbers can be as high as 4,100 (in Gigantiops) While ant reliance on visual cues varies across species, all
and 3,200 (Myrmecia). Additionally, several species have ants rely heavily on chemical communication and use their
ocelli on their forehead. These are light-sensitive eyespots antennae as their main sensory organs. The antennae are
that likely have a role in navigation. Ocelli are usually absent typically elbowed, consisting of 4 to 12 individual shorter
in workers and more commonly seen in queens and males. segments called antennomeres. The base of the antenna is
Ants are not primarily visual insects, and in ant brains the one long, single segment—the scape. This is followed by the
optic lobes are much smaller than are seen in close relatives funiculus, which is made up of multiple shorter segments.
such as wasps and bees. This may be because most ants do Segments in the funiculus may be of equal size or gradually
not fly and those that do spend only a small part of their life increase in width. Occasionally a few of the end segments
flying. Ants spend less time relying on visual sensory cues form a bulbous club. The scape and funiculus form an
and are more heavily reliant on other senses such as elbowed joint, a unique feature of the ants. The antennae
nestle within a socket on the front of the head. Some species
of ants have antennal scrobes—grooves on the sides of the
head that the antennae can be retracted into.

Antennae are moved around by muscles inside the head as


well as in the base of the antennae. These muscles are
controlled by the dorsal lobe in the brain, which receives
mechanical sensory information input from the antennae.
Olfactory receptor neurons in the antennae are connected to
the antennal lobe. Antennal lobes are large in ants relative to
other insects because they rely more on smell than sight and
because they use pheromones for communication. Antennal
lobes can process ordinary smells as well as pheromones.

L E F T: Based on the number of ommatidia, Gigantiops


destructor has the largest eyes of any ant, which it uses to
locate prey while foraging in the forests of South America.

24 WHAT IS AN ANT?
MO UTH PA RTS
Ants have biting and chewing mouthparts, often with
distinctively shaped and well-developed mandibles. Ant
mouthparts have an upper lip (labrum) and lower lip (labium).
They also have a pair of mandibles and maxillae. The maxillae
and labium each have sensory palps that can feel and smell
food as it is being eaten. Mandibles are typically triangular
in shape, with a smooth outer edge and a toothed inner
masticatory margin. However, mandibles have evolved into
a vast array of shapes and sizes, from toothless to double-
forked teeth on the masticatory margin, to overall shapes
that are variously short or elongated, straight or curved.

Mandibles are used by ants for a variety of tasks, including


feeding, attacking, and defending, as well as handling and
manipulating objects. Mandibles are used for excavation
and construction of nests. Carrying behavior in ants requires
a specialized skeletomuscular modification of the head and
neck muscles. Some ants are known to carry prey between 30
to 90 times their own weight and perform delicate controlled
head movements while carrying these objects. As well as food
items, ants will also carry liquids suspended as a drop
between their mandibles. Worker ants will carry brood (eggs, ABOVE: The trap-jaw mandibles
larvae, and pupae) to move them around different parts of the A B O V E L E F T: Located above the of Myrmoteras iriodum have 12
nest or to a new nest location. This requires a sensitive touch eyes, the antennal scrobes of teeth and are usually held open
using mandibles in a careful and deliberate way so as not to Mycetomoellerius tucumanus are at 280 degrees. These, and large
damage the soft-bodied early life stages of the colony. Finally, visible as deep channels on the compound eyes, make these ants
ants are even known to carry each other. top of the head. efficient hunters of small prey.

SENSORY STRUCTURES 25
I N T E R N A L A N ATOMY

N ERVES discontinuous gas exchange, where there is a cyclical pattern


In the ant nervous system, bundles of nerve cells form created through the opening and closing of spiracles at
ganglia that are connected by nerve cords to create the central different times, meaning that oxygen is taken in during
nervous system. There is essentially one long nerve cord activity and carbon dioxide is released during rest periods.
running from the brain through to the abdomen with The respiration system of ants delivers oxygen directly to
branching nerves along it. Ant brains are like most insect tissues, reducing the requirement for a circulatory system to
brains, containing different regions that relate to various, perform gas exchange. Ants have an open circulatory system
often sensory, functions. The optic lobes are the visual with a single long tube running the length of the body that
centers, comprising the lamina, medulla, and lobula, while is equivalent to the heart. A pulsing movement moves
the antennal lobe is the primary olfactory center. There are hemolymph around the ant’s body. Hemolymph is the ant
also protocerebral lobes, the central body, and the mushroom equivalent of blood; it moves nutrients, hormones, and waste
body (made up of the calyx and the mushroom body lobes). around the body while also circulating hemocytes—cells
Fused to the back of the brain is the subesophageal ganglion. responsible for insect immunity.
This bundle of nerves branches into the mouthparts and acts
in sensory detection and muscle movement. These nerves also D I G ESTI O N
control the head and neck movements. Following on from the Ants follow the same basic digestive system plan as most
subesophageal ganglion is the first thoracic ganglion. This insects in that they possess a fore-, mid-, and hindgut. The
then forms a chain of ganglia that run through the body as foregut is the most complex, containing the oral cavity,
thoracic, petiolar, and gaster ganglia, although the numbers followed by the pharynx, esophagus, crop, and proventriculus.
of ganglia are not well known in many ant species. The The esophagus is a long tube that moves food through the
mushroom body may be involved in cognitive functions such insect body to the crop, a ball-shaped sac that acts as a food
as learning and memory. The role of the central body in ants storage area and is located within the gaster. In ants the crop,
is unknown but may be related to organizing behavior, for often referred to as the “social stomach,” is used by foraging
example in polarization vision or leg control. On average, worker ants to store liquid foods such as nectar or insect
larger ants have larger brains, with males having smaller honeydew. In a process known as trophallaxis, this food is
brains than female queens and workers. then regurgitated and shared out among other members of
the colony. The proventriculus is a muscular valve that
C I RCU L ATI O N A N D R ESPI R ATI O N facilitates the process of trophallaxis. Any food allowed to
While an ant’s respiration occurs in metabolically active pass into the midgut is digested by the ant, but food held in
tissue, the process of gas exchange (receiving oxygen and the crop due to the constriction of the proventriculus is part
removing carbon dioxide) occurs through spiracles—tiny of the food supply for the colony’s social stomach. The
holes in the exoskeleton. Spiracles are connected to trachea, proventriculus also acts as a separation point to prevent
and these are connected to even finer branching tracheoles digestive enzymes from the midgut flowing into the crop.
that perform gas exchange with cells throughout the body. Digestion occurs in the midgut, where absorption of
Gas exchange can occur by diffusion of the gases through the nutrients takes place. The hindgut function is devoted to
tracheal system, where a continuous movement of oxygen the process of reabsorption of water and salts.
and carbon dioxide occurs. Some ant species show

26 WHAT IS AN ANT?
I N TER N A L BO DY SYS TE MS

Crop Midgut

Rectum

Brain

Pharynx

ABOVE: The internal organs of ants include KEY


the brain and nervous system, the circulatory Digestive system Circulatory system Nervous system
system, and the digestive system.

INTERNAL ANATOMY 27
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BRINGS-DOWN-THE-SUN TELLS ABOUT HIS
FATHER

When Brings-Down-the-Sun came to our camp the following evening,


he took a seat by the fire. For a while he meditated, smoking his
everyday pipe in silence, and then he began:

“There is a trail we call ‘The Old North Trail.’ It runs along the Rocky
Mountains outside the foothills. It is so old no one knows how long it
was used. The horse trail and travois tracks were worn deep into the
ground by many generations of Indians.

“My father told me that this old trail was started ages ago by an Indian
tribe coming down from the north; and other tribes followed in their
tracks. I have followed the Old North Trail so often I know every
mountain, stream, and river of its course. It ran from the Barren Lands
in the north to the south country, where people have dark skins and
long hair over their faces (Mexico).”
BRINGS-DOWN-THE-SUN AND THE AUTHOR

By this time it was growing dark and the fire burned low. No one spoke
until the silence was broken by the mournful howling of a wolf in the
near-by hills. Then Brings-Down-the-Sun continued:

“The wolf is our friend and we do him no harm. The Indians have a
saying, ‘The gun that shoots a wolf or coyote will never again shoot
straight.’

“I never heard of a wolf that did not wander. They raise their young in
one place and then move on to another. They like to run all over the
country.

“My father’s first name was Running Wolf. His father, Little Mountain,
gave him that name, because a wolf appeared in a dream and advised
him, saying:

“ ‘I am the head-chief of all the wolves and my name is [211]Running


Wolf. You often hear my voice; my tracks are everywhere and I shall
always continue to wander. Name one of your sons after me; and if he
has a son, let that name be handed down; all of your descendants who
bear my name will have long life and good luck.’

“I have now the wolf-nature, because my father bestowed the name of


Running Wolf upon me. Like the wolf I wander over the plains and
through the mountains; I never like to stay long in one place.

“Now I am going to tell you more about my father—how my


grandfather, Little Mountain, happened to honor him with the name of
Running Wolf, although he had two older brothers.

“One day in early summer, when my father was only a boy, he saw a
band of warriors gather near my grandfather’s lodge; he was then
head-chief of the tribe. They were dressed for war and had their
horses painted with war signs. They stood in a circle, holding a big
rawhide between them, upon which they beat with sticks like a drum
and sang a wolf song. They marched singing through the camp,
saying farewell to friends and relatives; and then started south on a
war expedition.

“Now this was very exciting to my father; his two older brothers were
among the warriors. He wanted to go too. But he knew they would not
take him because he was too young.

“After the expedition had gone, the boy ran into the lodge and took a
bow and arrows. He told his father, the head-chief, he was going for a
hunt; but he rode fast and overtook the war party. They tried to send
him back. But one of his brothers said:
“ ‘If he is so eager, let him come; he can look after one of the pack
horses.’ Thus it happened that my father went with that war party.

“One night, after they crossed the Yellowstone River, [212]the boy was
wakened by a noise. He scouted around and saw a band of hostile
Indians coming into camp. They were some of the Snakes and he
gave the alarm. The Blackfoot warriors made ready to attack, but
waited until just before dawn.

“In the fight my father made a wonderful shot and killed a Snake
Indian. It was the only scalp they took on that trip. When the warriors
came back to the Blackfoot camp, they waited on the summit of a hill,
until a big crowd came out to meet them.

“Then they made known to the people how the boy was the only one
to kill an enemy. They showed the Snake scalp and the head-chief
was proud of his son. He tied that scalp to a long pole and told the boy
to hold it aloft, and to shout as they rode triumphantly through the
camp: ‘My name is Running Wolf; I am the youngest of the war party
and the only one to kill an enemy. Behold! Here is the scalp.’ Thus it
happened that my father got the name of Running Wolf.

“When my father was head-chief of the tribe, he went by the name of


Iron Shirt, because he was accustomed to wear a shirt which was
decorated with shining pieces of metal. He was also head man of the
band of Grease Melters. He was a large and muscular man, with fine
mind and a wonderful memory. He knew all the legends and lore of his
tribe. He could tell the age of a horse by its whinny and of a man by
the sound of his voice. He kept ‘winter-counts’ by making pictures on
buffalo robes. He recorded important events in the history of the tribe
—places of tribal camps, battles and the names of war chiefs, years of
smallpox, summers of drought and winters when snows were deep
and food scarce. He kept count of the winter when many of our people
died from the cough-sickness, the winter when the children broke
through the ice, when some moose came into our camp, also the
winter when we had to eat dogs to keep from starving, the time a
[213]herd of antelope broke through the ice, when we caught some
antelope in the deep snow, when buffalo were scarce, and the time we
made the first treaty with white men.

“Sixty-one winters have passed since we had our first great sickness
of smallpox (1836); forty-two, since we had the big camp on the
Yellowstone River (1855), the time eight Indian tribes came together
and our head-chiefs were Little Dog, Big Snake, and Lame Bull;
twenty-seven winters since the coming of the North-West Mounted
Police (1870), and twenty-one since the bad winter, when many of our
horses were frozen to death (1876).

“I was born in the spring, the year the first white men appeared in our
country. And I was still a young boy when my father became the owner
of his first Medicine Pipe. This happened in one of our tribal camps,
which was being held in mid-summer. Wolf Child was the owner of a
Pipe and chose my father as his successor. He told the medicine pipe
men he wanted them to take my father.

“Now it happened that my father was a ‘bear-man’—that is, his power


came from the grizzly bear. He had a sacred bearskin inside his lodge,
which he kept hanging from the lodge-poles, just over his couch. The
word ‘bear’ must never be spoken in the presence of a Medicine Pipe;
it has an evil influence. For this reason the medicine pipe men were
always afraid to offer a Pipe to my father, with the sacred bearskin so
near.

“But Wolf Child, the owner of this Pipe, advised there was no danger
from the skin. He said my father had great power; besides it was
possible for them to drive away the evil by burning sweet pine as
incense.

“Thus Wolf Child persuaded his friends and overcame their fears. It
was after midnight, when I heard the medicine pipe men stealthily
enter our lodge. Wolf Child came first, with the sacred Pipe hidden
under his robe. They caught [214]my father asleep; and, when he woke
up, they offered him the Pipe. He took it in both hands and they all
began to drum and sing. After that my father smoked the Pipe and
said:

“ ‘I am the owner of many horses, which of them do you want?’

“Wolf Child answered: ‘Your black buffalo-horse.’ Now this was the
most valued horse in my father’s herds. He was a famous race horse,
the fastest in the tribe; he was so high-spirited it took three rawhide
bridles to hold him.

“But my father did not hesitate. He answered quickly: ‘Take him, he is


yours.’ So Wolf Child got the horse and my father became the owner
of a Medicine Pipe. It was an honor to be chosen, although the Pipe
was a great burden. But my father could not refuse; no one ever dares
to turn down a Medicine Pipe. I know of an Indian who tried it, because
he did not want to give up a valuable horse. Misfortune came upon
him. His father-in-law died, then the horse; and finally the man died
himself. All because he refused to take over a Medicine Pipe when it
was offered to him.”

Origin of his Father’s Thunder Pipe

“Once I was camped with my father and grandfather on the St. Mary’s River.
We were near the mountains after beaver, which were plentiful then. One
day my father went alone on a hunt, following the trail of some elk to Chief
Mountain. At timber-line he came upon a band of mountain sheep and
trailed them towards the summit. Near the top of the mountain, he came
upon bad-smelling smoke coming from a deep hole. Into it he rolled a stone.
He waited to hear it fall. But no sound came back; only a cloud of smoke so
dense he could hardly breathe. Then he saw a thunder cloud coming down
the mountain and started to run. There came a crash and he fell to the
ground. A woman stood over him; her face was [215]painted black and she
had red zigzag streaks for lightning below her eyes. Behind her stood a man
with a huge weapon. My father heard him say:
“ ‘I told you to kill him quickly, but you stand there and pity him.’

“He heard the woman chant: ‘When it rains, the sound of the Thunder is my
medicine.’

“Then the man sang and fired his big weapon; it sounded like the crashing of
thunder, and my father saw lightning coming from the hole in the mountain.
Suddenly he found himself inside a cave; he could not speak, neither could
he raise his head. He heard a voice say:

“ ‘This is the person who threw the stone into our fireplace.’

“He heard some one beating a drum; and after the fourth beating, he was
able to sit up and look around. He was in the home of the Thunder. He saw
the Thunder Chief in the form of a huge bird, with his wife and children
around him. They all had drums painted with the claws of the Thunder Bird,
and its beak from which came streaks of lightning.

“Whenever the Thunder Chief smoked his Pipe, he blew two whiffs towards
the sky, then two to the earth; and after each whiff the thunder crashed.
Finally the Thunder Chief said to my father:

“ ‘I am the Thunder Maker and my name is “Many Drums.” You have seen
my great power and can now go in safety. As soon as you return to your
camp, make a Pipe just like the one you see me smoking. When you hear
the first thunder in the spring, you will know I have come from my cave.
Then it is time for you to take out your Pipe and hold it up. If you are ever in
a bad thunderstorm and feel afraid, pray to me saying:

“ ‘Pity me! Many Drums, for the sake of your youngest child,’ and no harm
will come to you. [216]

“Thus it happened that my father became the owner of a Thunder Pipe.


When he knew he was going to die, he gave this Pipe into my care. He said
it was a ‘long-time-pipe’ and must not be buried with him. I still have the
Thunder Pipe and smoke it only on important occasions.

“My father was skilled in the catching of eagles. He taught me how to take
eagles alive, and for many years the catching of eagles has helped to
support my family. But it was a hard and dangerous calling. I had to go to a
solitary place near the foot of the mountains and dig a hole in the ground
deep enough for me to stand in. I killed a coyote and stretched the hide on
sticks, laying raw meat along the sides, to look as if it were freshly killed. I
entered the pit before daylight, in order that no eagle could see; and covered
it over with branches and leaves. The coyote bait lay on top, just over my
head. I stood in that pit all day without food or drink. I could not even smoke,
lest the eagles might get the scent. Throughout the day I chanted the coyote
song, ‘I want the eagles to eat my body,’ because it had the power to attract
eagles to my bait.

“The Long Tails (magpies) generally came first. They walked around the
meat, chattering and saying to each other over and over, ‘Long Tails go
ahead and hang your sack upon a tree.’

“After a while an eagle would see the magpies eating the bait and come
near. At first he would be suspicious. When he walked upon the blind and
started to eat, I thrust both hands through the branches and seized him by
the legs. I drew him quickly into the pit and killed him by breaking his neck
with my foot, so that the wings fell to both sides and the feathers were not
injured.

“Golden eagles were the most profitable to catch; the Indians wanted them
because of their white tail feathers with black tips which they used for
headdresses and sacred bundles. [217]Bald eagles were scarce and hard to
catch. Some of them were so powerful they almost dragged me from the pit.

“In those days eagle-catching was a dangerous occupation, because of


grizzly bears. I remember an Indian, who held fast to his bait when a big
grizzly started to drag it away. The bear pulled off the branches and saw the
man in the pit. He pulled him out and tore him to pieces. When his relatives
came, there was nothing left but his bones.

“My father did not die in battle, nor of sickness, but of old age. After his
death, I became interested in religious things and came north to live. One
night I slept alone on a high hill of the prairies. I had a strong dream. The
Sun God came to me and said:

“ ‘My son, be not afraid. I give you my power and will guard you through life.’

“I took an interest in the Sun Dance and became one of its leaders. From
that time, I was no longer called Running Wolf. People called me Brings-
Down-the-Sun (Natósin-népe-e), because I had the power of the Sun.

“I have nine children living, four sons and five daughters. The names of my
sons are, Running Wolf, Iron Shirt, Double Walker, and Three Eagles. The
girls are, Long Hair, Turns-Back-the-Herd-Alone, Good Kill, Double-Gun-
Woman, and Whistling-All-Night.

“Towards the north lies the highest summit of the Porcupine Hills. No trail
leads to it and it is surrounded by a dense forest. The top is steep and is
covered with stunted pines. From that direction come our hardest storms.
When my oldest son died I went to that lonely summit. I did not want to see
any one. I stayed there night and day and fasted. Then I had a dream. The
Spirit of the Mountain came to me and gave me a Medicine Robe; 1 and with
it went supernatural [218]power—power to heal the sick. This wonderful
Robe had many skins of birds and wild animals attached. There were marks
to represent the Sun and Morning Star, also the constellations of the Bunch
Stars and the Seven Persons. I wore it in the ceremony of the Sun Dance,
when I stood before the people; and it gave me power to doctor the sick.

“That Spirit of the Mountain warned me to give up using the sweat-lodge,


lest more of my children die; and to wash daily in the river. Since that time I
have always bathed in the river every morning, even in winter when I have
to break the ice. I tell my sons to bathe regularly; and after they finish, the
women go in. I believe sickness can be warded off by keeping the body
clean and using sweet smoke for incense.”

[219]

This Medicine Robe of Brings-Down-the-Sun is now in the Blackfoot Collection,


1
American Museum of Natural History, New York City. ↑
[Contents]
CHAPTER XXXI
BRINGS-DOWN-THE-SUN TELLS ABOUT MEN’S
SOCIETIES

The Braves

“I am a member of the Society of Braves. It was started long ago by an


Indian. He had a strong dream in which he saw a band of dogs and the
way they acted. The Braves ruled the camp and helped our chiefs to keep
order. We punished men and women who quarreled; we sometimes killed
people who disobeyed our orders.

“When it was time for our tribe to move camp, we marched with the
beating of drums and singing, each member carrying a knife and a bow
and quiver full of arrows. We went to the center of camp and spent the
night curled up on the ground like dogs. The day our tribe moved, we
stayed behind and acted like dogs, eating all the food that was left; like
dogs we followed slowly and entered camp after all the lodges were
pitched.

“We placed our big lodge in the center of camp, taking possession early,
on the morning we danced. Our leader wore a coyote skin for a headdress
with the tail hanging down behind. He was called Wolf-Skin-Man and
carried a short lance, which was decorated with feathers. The next in rank
carried a willow branch. He wore a robe with buffalo hoofs attached, which
rattled when he danced.

“On the left of our leader sat the ‘white braves,’ with white painted lances
stuck into the ground in front of their seats. Their bodies were painted
white; and they had yellow stripes across nose and eyes. Two other
members called ‘water braves’ were painted black; they carried bladders
on [220]their backs for water pails and bags for back-fat and pemmican.
There were also ‘black braves’ carrying black lances, with bodies painted
black and black stripes across their faces; and ‘red braves’ with red lances
and red painted bodies.
“Two other braves carried bows and arrows instead of lances. They had
their faces painted to impress the spectators; their bodies were covered
with red and on their faces was the bear sign—black streaks down over
the eyes and at each corner of the mouth. They wore their front hair short
and made it stand straight up by covering it with paint. They had fringed
shirts made from the smoked tops of old lodge covers, belts of bearskin
and arm-bands of bearskin with bear claws attached; for headdresses
they had strips of skin with bear ears and two claws attached to look like
double ears.

“Whenever we danced outside our society lodge, we sat in an open circle


with four drummers in the center, our lances stuck into the ground with
points down. Wolf-Skin-Man as leader was at the head of our circle,
opposite the opening, with the two bear braves outside. They covered
themselves with robes, as if they were bears lying in a den.

“Our leader, Wolf-Skin-Man, danced first, blowing his bone whistle as a


signal for the others to follow. Then we dropped our robes and followed
him, blowing whistles and bearing our lances. We danced slowly in a
circle, leaning forward and holding our lances near the ground and acting
like dogs looking for places to lie down. The white-painted braves drove
the others before them with their lances, but stopped as soon as the two
water braves appeared. Then came the brave with the willow branch, who
could not stop dancing until the two grizzly bears appeared.

“The bears only danced when they felt like it. They lay in their den and did
as they pleased. Sometimes the spectators threw things at them to make
them dance. When they [221]were ready they got up slowly, holding their
hands the way bears do their paws. They danced leaning over, hopping
along in short jumps with their feet together, as bears are accustomed to
do. They aimed with bows and arrows, and drove the other dancers back
to their seats.

“When we wanted to bring our dance to a close, the two bears pretended
they were going to shoot at the spectators with sharp-pointed arrows; but
they changed quickly to painted arrows without points and shot them over
the heads of the people. Then we ran over the prairie, in the direction the
arrows flew, taking off our moccasins and throwing them away.

“We marched through camp singing our society song and shouted our
orders to the people. It was our custom to take anything we wanted, even
food from the kettle, as dogs do. We stopped to dance at the lodges of
prominent chiefs, who were expected to give us presents of food and
clothes. If any people bothered us or held us back, the bears shot at them
with their arrows.

“If our leader told us to allow no one to chase buffalo, then we were the
watchmen of the herds. If the buffalo were frightened and ran away,
scared by some one who chased secretly, we followed that man and took
away his horse and weapons; we tore off his clothes and whipped him; we
sent him back to camp naked and on foot. We alone could look for buffalo;
and, when we found plenty of them, we moved the tribe that way. We also
punished women who picked berries against our orders by tearing their
lodges to pieces.

“Men who joined the Society of Braves had to face danger whenever it
came; they could not turn away. When we decided it was time to change
our camping place, we made a feast in our society lodge and invited the
head-chief. After we had eaten and were smoking together, our head-chief
would say: [222]

“ ‘My children, why have you asked me to come here? What is it that you
want’?

“And our leader would answer: ‘Grass is scarce for the horses; the water
is poor and the ground no longer clean; it is time for us to move to another
place.’

“Then the head-chief would reply: ‘In the morning we shall break camp; it
is too late to-day. Tell the people to bring in their horses from the hills and
to picket them close to their lodges; we shall start at the rising of the sun.’ ”

The Society of Mosquitoes

“This society was formed many years ago by an Indian who hunted in a
place where there were great numbers of mosquitoes. They came in
swarms and bit him all over; he lay on the ground and lost all feeling; he
heard strange voices singing:

“ ‘Mosquitoes, mosquitoes, get together, get together.


Mosquitoes, get together,
Our friend is nearly dead.’

“And then he saw mosquitoes beginning to dance. Some were red and
others yellow. They had claws attached to their wrists and long plumes
hanging from their hair. They sat in a circle and sang; they jumped up and
down, springing this way and that, always dancing in the direction of the
sun. He heard a voice say:

“ ‘Brother, because you were generous and let us drink freely from your
body, we give you our society of mosquitoes; we make you the leader.’

“Then that man came safely home and started the Society of Mosquitoes.
Its members wore buffalo robes with the hair side out. Some painted
themselves red and others yellow, with stripes across nose and eyes.
They wore plumes in their hair and eagle claws attached to their wrists to
represent the bills of mosquitoes. [223]

“When the mosquito society gave their dance, they sat in a circle around
the drummers; the leader was at the head, with yellow mosquitoes on both
sides. After each song, they held down their heads and made a buzzing
noise in imitation of mosquitoes. After repeating this dance four times,
they scattered and went through the camp. Any person they met, they
scratched with their eagle-claws, and said: ‘Now I shall take blood from
you.’

“Any one who resisted or tried to run away, they caught and scratched
hard. But those who offered themselves freely, and invited them to take
their fill, they did not hurt. Because, if a mosquito is left alone, its bite does
no harm.”

The Society of Kit Foxes

“This was one of the oldest of all our societies. It was started by an Indian
named Elk Tongue, who dreamed about a kit fox. It invited him into its den.
And there he saw the chief of the foxes with many foxes seated around
him. Before he left their den, the fox chief said:

“ ‘When you return to your tribe, take a foxskin for your medicine; wear it
always on your back and my power will go with you. Form a Society of Kit
Foxes. Gather together some young men and show them how to dress
and to dance. Tell them, if they do these things and never kill any foxes,
they will have benefit. But it will be bad luck for them to harm a fox.’

“After Elk Tongue came home he started the Society of Kit Foxes. The
members had great power. The ceremony was secret and the people were
afraid; it was dangerous to talk about it. Elk Tongue was their leader. He
alone knew the secrets and told the other members what to do. Before he
died he taught the ceremony to his son and told him all the secrets. After
that the power of the foxskin was handed down from father to son through
many years; and the members [224]of that society taught their children
never to harm a fox.

“When the Kit Foxes gave their ceremony, they opened up two lodges and
made them into one. For four days and four nights they sat inside, painting
and dressing themselves, singing and making ready, only appearing
outside their dance lodge at night.

“But, on the fifth day, they came out and marched through the camp. Their
leader wore the foxskin with its head in front, the ears on top, and the skin
itself, with small bells attached to the tail, hanging down his back. He
carried a bow and arrows, which were painted green; his body was also
painted red, his face green, to look as frightful as possible and to make
people afraid.

“The second in rank, called the white-circle-man, carried a long lance, with
one end bent into the form of a hook; it was covered with white swan’s-
down and had white eagle feathers attached at regular intervals along its
staff.

“The next in rank had a long lance in the form of a hook, which was
wrapped with otter-skin and had black and red feathers attached. The
other members carried pointed lances decorated with feathers, and small
pipes painted red. They all painted their faces and wore eagle feathers in
their back hair. Around their legs were wide bands of otter-skin with small
bells attached.

“When the Kit Foxes marched through the camp, they formed in the shape
of a fox head. The chief went first to represent the nose; behind him were
the second and third men for the eyes; then came the other members in a
group, all together representing the head of a fox. The two second men,
as the eyes, watched the chief who was the nose, or leader; they acted as
he directed and the rest followed after.

“For their dance, they sat in lines, with the regular members in the first
line. If any were withdrawing from the society[225]—giving up their lances
to new candidates—they sat in a second line; and the wives of the
members sat behind.

“As soon as the drums began, the leader started the dance; and the two
circle-men with white lances followed. After them came the other members
with plain spears. They danced in pairs, the way Kit Foxes run together.
They gave short, even jumps with feet close together, imitating the
movements of a fox. They barked and moved about, first in one direction,
then in another, just as a fox is accustomed to do. The two circle-men
(eyes), barking and swinging their spears, danced between the two lines.
They did not move in a straight line, because a fox never goes straight; his
tail always seems to guide him. When the white-circle-men shouted, ‘It is
enough,’ the dance stopped and the members returned to their seats. But,
after a short rest, they began again. And that is what I know about our
societies.”

[226]
[Contents]
CHAPTER XXXII
BRINGS-DOWN-THE-SUN TELLS ABOUT THE
BIRDS AND THE STARS

We were seated by our camp-fire with the old chief and his family.
My Indian sister, Strikes-on-Both-Sides, and Long Hair delighted the
children, making whistles from cottonwood bark and toy lodges from
leaves of the balsam poplar, winding them around their fingers into
the shape of little tepees and fastening them together with twigs.
They set them up like real lodges and made them into a big camp in
the form of a circle.

Then we gave Brings-Down-the-Sun the collection of roots and


herbs we gathered especially for him; and a rare medicinal root
called “sharp vine” by the Indians, one of his favorite remedies for
breaks and sprains. He was so glad to get it that he chanted and
prayed over it. After putting our present carefully in an old medicine
sack, he seated himself apart from the others, and with dignity
waited for the women and children to become quiet.

From his manner I knew he was ready for his Indian stories and
hastened to prepare notebooks and writing equipment. The old chief
was watching, for he said with a smile:

“My white son there reminds me of a squirrel; he runs in one


direction as if to steal something, then darts quickly in another; he
never sits still a moment; he is always on the move.”

At that moment a woodpecker with red-crested head called from a


tree top. Brings-Down-the-Sun pointed to the bird and said: “He calls
to the worms and bugs to stick out their heads; he is hungry and
wants to eat them.” [227]

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