Latin american insects entomology c hogue (university of california, 1993)

LATIN AMERICAN INSECTS AND ENTOMOLOGY Charles L... LATIN AMERICAN INSECTS AND ENTOMOLOGY Charles L... T h e other topics of discussion, such as ecology and the study of insects, have

LATIN AMERICAN INSECTS AND ENTOMOLOGY

Charles L Hogue

5>*il^Vc - C ,A ' ' ■ > ' ■

Xfti

Lantern bugs and cicadas, from Madam Merian's Metamorphoses insectorum surinamensium

(1705) Note the imagined specimen at the bottom of the plate which combines features of the two

insects

LATIN AMERICAN INSECTS AND

ENTOMOLOGY

Charles L Hogue

UNIVERSITY OF CALIFORNIA PRESS • Berkeley • Los Angeles • Oxford

!

University of California Press

Berkeley and Los Angeles, California

University of California Press

Oxford, England

Copyright © 1993 by The Regents of the University

of California

Library of Congress Cataloging-in-Publication Data

Hogue, Charles Leonard

Latin American insects and entomology / by Charles L Hogue

CIP Printed in the United States of America

1 2 3 4 5 6 7 8 9

The paper used in this publication meets the mini­

mum requirements of American National Standard

for Information Sciences—Permanence of Paper for

Printed Library Materials, ANSI Z39.48-1984 ©

care e n s u r e d t h a t Latin American Insects and En­

tomology, C h a r l e s H o g u e ' s last major w o r k ,

would a p p e a r w h e n a n d as his f a t h e r w o u l d have wished

The ubiquity of spirits and the impossibility of killing them seem to personify a feeling of helplessness in the face of an environment so beautiful and so cruel On the river or work­ing in a garden the sun hurts, "It is eating," the Sharanahua say, and heads ache for the rest of the day T h e incessant gnats feed all day, and, as one lies in a hammock, someone leans over and slaps hard and says, "sandfly," and a black fly, fat with human blood, falls dead Sundown is a moment of relief which even a hundred mosquitoes cannot mar

In the forest someone shouts to warn of an

uula, the huge stinging ants that make one

drunk with pain, and, reaching for a

handhold on a tree, one must avoid a swarm

of red fire ants Returning, one looks for ticks, huge tapir ticks, gray and voracious, or worse, the almost invisible tiny red ticks that burrow into the skin and hurt for a week The women dig the egg sacs of chiggers out

of toes skillfully so that the sac does not break

to leave a budding worm to swell the foot, and they break each and every tiny egg with a needle so that it does not lie in wait for an­other bare foot An infected gnat drops a worm's egg into the leg while sucking blood, and two weeks later the pain of the worm turning in the leg is excruciating, and it must

be removed by daubing an old, foul, drop of tobacco juice on the skin and slowly winding the worm out on a stick Women and girls pick lice out of men's hair and their own, break them in their teeth and eat them When faced by a new animal or insect, I learned to ask both, "Do we eat it?" and "Does it eat us?"

Janet Siskind,

To Hunt in the Morning

10 MOTHS AND BUTTERFLIES 292

11 FLIES AND MIDGES 360

12 SAWFLIES, WASPS, ANTS, AND BEES 405

13 INSECT STUDY 470 Included Insect and

Arthropod Taxa 499 Index 521

Preface

The idea for this book germinated in my

mind for many years after my christening

in Latin American insect research As a

result of travels to many countries, it

became acutely apparent to me that a

comprehensive entomological work was

sorely needed to serve the many people,

both visitors and residents, interested in

insects and their other terrestrial arthropod

relatives The curiosity of tourists and

general natural historians needed satisfac­

tion, and the more serious minded student

and practicing professional needed an

up-to-date review of the subject After a long

period of note taking and preliminary

organization of my then chaotic knowledge

of the subject, I resolved to fashion such a

volume

Some of my colleagues were incredulous

that I could cover such a vast territory

But my experience writing general insect

guides told me that with cautious choosing,

I could make something really useful,

though, of course, far from complete In

fact, writing this book has been an exercise

in selectivity, especially with respect to the

choice of taxonomic groups to include I

relied on my own experience and the

experience of others in this process, and I

have tried to give information on the most

common, conspicuous, or otherwise nota­

ble (economically or historically important)

units, whether species, genera, or higher

groups T h e other topics of discussion,

such as ecology and the study of insects,

have also been presented with an eye to the reader's need for an overall understanding

of what has transpired and is transpiring in Latin American entomology and to provid­ ing a framework for review and citation of pertinent literature

Some who read this book will feel slighted because of the lack of coverage of topics of particular interest to them, or they may consider that important facts, taxa, or publications have been omitted I can only ask these readers to remember the vastness

of the subject and the necessity for extreme conservatism in choices of matter for inclu­ sion Of course, I welcome suggestions for additions or changes in emphasis for future editions

I have designed the book to answer questions In my language and in the selection of taxa/phenomena, and points of information about each, I have been guided

by my perception of what most readers want to know rather than a desire to produce an encylopedia of all the facts that might be recorded The technical litera­ ture, to which I have so freely referred, will serve the latter purpose Indeed, to present in-depth data, keys to identification, and exhaustive treatments of even the major categories of Latin American insect life would require many volumes and years of effort to produce and would not produce the ready, readable, and portable text that I think is most needed now

Acknowledgments

For reviewing sections and offering criti­

cisms and information on many topics, I

have been fortunate to have the expert

assistance of many specialists I am greatly

indebted to the individuals named below

for information, identifications, and count­

less improvements in the manuscript (the

appearance of their names does not neces­

sarily imply agreement with my final inter­

pretations and statements): J Richard An­

drews (Náhuatl names), Arthur G Appel

(cockroaches), Phillip A Adams

(Neu-roptera), Richard W Baumann

(Plecop-tera), Vitor Becker (Lepidop(Plecop-tera), Jackie

Belwood (Orthoptera), Harry Brailovsky

(bugs), A Brindle (Dermaptera), Jacob

Brodzinsky (fossils), Lincoln P Brower (but­

terflies), John C Brown (Lepidoptera),

Richard C Brusca (Crustacea, arthropod

anatomy), Gary R Buckingham (useful

insects), James L Castner (Orthoptera),

Gilbert L Challet (aquatic beetles), John A

Chemsak (Cerambycidae), James C

Coken-dolpher (harvestmen), Julian P Donahue

(Lepidoptera), John T Doyen (darkling

beetles), Richard D Durtsche (study),

W D Edmonds (dung scarabs), George F

Edmunds, Jr (mayflies), K C Emerson

(Mallophaga), Marc E Epstein (Lepidop­

tera), Terry L Erwin (beetles), Arthur V

Evans (beetles), Eric M Fischer (Diptera),

Oliver S Flint, Jr (Trichoptera), Will Flow­

ers (aquatics), Manfredo A Fritz

(Sphe-coidea), Richard C Froeschner

(Hemip-tera), David G Furth (ecology), Rosser W

Garrison (Odonata), P Genty (palm pol­

linators), Edmund Giesbert (Cerambyci­

dae), Eric E Grissel (Chalcidoidea), Robert

J Gustafson (botany), Alan R Hardy abaeidae), Brian R Harris (butterflies), Steven Hartman (mantids), Henry A Hes- penheide (beetles), Frank T Hovore (Cer­ ambycidae), Chistopher A Ishida (prac­ tical entomology), D K McE Kevan, deceased (orthopteroids), James E Keirans (ticks), John E Lattke (general), Claude Lemaire (Saturniidae), Herbert W Levi (spiders), James E Lloyd (Lampyridae), Wilson R Lourenco (scorpions), Richard

(Scar-B Loomis, deceased (chiggers), Robert J Lyon (gall wasps), Volker Mahnert (pseu- doscorpions), Sergio Martinez (fossils), Mildred E Mathias (botany), Eustorgio Méndez (medical entomology, parasites), Arnold S Menke (Cynipidae and other Hymenoptera), Edward L Mockford (pso- cids), Jacqueline Y Miller (Castniidae), Michael J Nelson (medical entomology), David A Nickel (orthopteroids), M W Nielson (leafhoppers), Lois B O'Brien (Ho- moptera), David L Pearson (tiger beetles), Stewart B Peck (cave insects), Norman D Penny (Neuroptera and general), Don R Perry (canopy insects), Manuel L Pescador (mayflies), John T Polhemus (aquatic Hemiptera), Diomedes Quintero Arias, Jr (whip scorpions), Shivaji Ramalingam (mos­ quitoes), Edward S Ross (Embiidina), H F Rowell (Acrididae), Raymond E Ryckman (ectoparasitic bugs), Ann L Rypstra (spi­ ders), Jorge A Santiago-Blay (scorpions), Jack Schuster (Passalidae), Terry N Seeno (beetles), Rowland M Shelley (myriapods), David R Smith (Symphyta), Roy R Snell-

ing (Hymenoptera), Omelio Sosa, Jr (his­

tory), Paul J Spangler (aquatics), Lionel

A Stange (Neuroptera), Orley R Taylor

(honeybee), Donald B Thomas

(heterop-terans and veterinary entomology), Carlos

Trenary (history), Fred S Truxal

(No-tonectidae), Alan Watson (Lepidoptera),

Thomas K Wood (Membracidae), A

Will-ink (history and study), Stephen L Wood

(Scolytidae), and Thomas J Zavortink

(mosquitoes)

For suggestions and information on

diverse or multiple topics, I also have many

other entomologists to thank, including,

José Artigas, Tomás Cekalovic, Ana Lia

Estévez, Carlos H W Flechtmann, Luis F

Jirón, Alberto and Beatriz Larraín, Carlos

Machado Allison, and Irene Rut-Wais

There are several colleagues who are

responsible for more pervasive involve­

ment in the work as a whole, who have

reviewed and contributed to the entire

manuscript, to whom I owe a special debt

of gratitude: Terry L Erwin, Arthur V

Evans, Gerardo Lamas, Jack Longino,

Scott E Miller, José Palacios-Vargas, Nel­

son Papavero, and Allen M Young For

tolerance of my many probings of their

knowledge and points of view, I am particu­

larly appreciative of the contributions of

Julian P Donahue, Chris Nagano, Roy R

Snelling, and Fred S Truxal

I am also indebted to the many local

friends and contacts I have made on my

Latin American travels—guides, foresters,

farmers, Native Americans, and many oth­

ers, too numerous to name, who freely

shared their firsthand field knowledge of insects and thus contributed to the original­

ity of this book in countless ways

To all who worked with the manuscript primarily in an editorial capacity, I wish to express gratitude, especially to Ernest Cal- lenbach Several librarians contributed in

no small way by finding and interpreting many difficult areas of the literature:

Nicole Bouché, Katharine E S Dona­

hue, Jennifer Edwards, Donald McNamee, Kathy Showers (Los Angeles County Mu­

seum of Natural History) and Nancy rod (University of California, Berkeley, Entomology Library) The preparation of the ink drawings were greatly assisted by Leland E Dietz and his Xerox machine

Axel-My appreciation is also due to Don Meyers for his careful and considerate handling of

my black-and-white photographic needs,

as well as to Tracy Robertson and James Robertson for other technical assistance with the figures

I thank James L Castner, George Dodge, and James N Hogue for allowing me to use those color photographs bearing their names in the captions

Finally, I acknowledge my wife, Bar­

bara, for her critical role in helping this task to completion, primarily her enor­

mous patience with my needs for time and other of our mutual resources She also typed most of my original drafts and did a great deal of editing For these contribu­

tions, not only extended here but lovingly provided in support of most of my entomo­

logical career, I dedicate this book to her

In discussing broad geographic regions,

I use the terms "Latin America," "New World tropics," and "Neotropics" (and their adjectival forms) as precisely as possible

The first refers to the most inclusive politi­

cal region within the bounds stated above;

the second refers to lowland, moist to wet, forested biotypes only; and the third refers

to the biogeographic region that includes South America, the West Indies, and tropi­

cal North America

In this volume, I discuss true insects and other kinds of terrestrial arthropods and related creatures (arachnids, millipedes, centipedes, onycophorans, etc.) commonly thought of as "insectlike." In many places,

it is overly cumbersome to be exact when referring to these groups, although I have tried to avoid misleading the reader by adding some explanatory phrase, such as

"and other terrestrial arthropods," but I have found it difficult to be perfectly consistent in doing so T h e meaning of a broader grouping is sometimes implied by the term "insect."

For each kind of insect or other ar­

thropod discussed in a separate section in

the systematic chapters, I open with a list of the applicable scientific names of the cate­ gory, including the most commonly used synonyms, and its place in the nomen- clatural hierarchy This is followed by estab­ lished vernacular names in Spanish, Portu­ guese, and other regional languages I use the widely familiar "Quechua" in place of the more correct "Runasimi." No attempt is made to give a complete synonymy, as this would require a lengthy treatment of its own The determination of plurals in some antique or indigenous languages is a prob­ lem, and some of those given may not find acceptance by purists I have replaced a few common names, for example, "wax bugs" for the order Homoptera, "dragon-headed

bugs" for Fulgora spp., and "big-legged

bugs" for the family Coreidae I feel that these are more appropriate and correct than previously used names and that they are useful to enhance common parlance about these often-mentioned taxa T h e r e are a few new common names (e.g., "viper

worms" for Hemeroplanes sp., Sphingidae;

"flag moths" for the subfamily Pericopinae

of the Arctiidae; and "shiny scarabs" for the subfamily Rutelinae of the Scarabaeidae The chapter on study of insects is in­ cluded as information for novices and to enhance the use of the book for teaching and for ready reference for professionals Much of the information has not been compiled elsewhere

Citations to the literature were chosen according to certain constraints First, they are always given as authority to, and source

of further information on, topics of special

1

interest, statistical s t a t e m e n t s , historical r e ­

m a r k s , a n d such I also i n c l u d e basic refer­

ences to g e n e r a l subjects o r sections o r

b r o a d categories b u t only t h e most m o d e r n ,

Because I u s e this f o r m a t , a few r e f e r e n c e s

have b e e n r e p e a t e d , obviating t h e n e e d for

cross listings a n d m a k i n g it easier for t h e

r e a d e r to find t h e m I h a v e seen all refer­

ences e x c e p t those r a r e o n e s followed by

"[Not seen.]." S o m e n e w observations a r e

ble; b u t generally, I follow v e r n a c u l a r

("wing covers") with technical s y n o n y m s

a r e based o n m u s e u m s p e c i m e n s entirely o r

in p a r t to confirm details p r e s e n t in existing illustrations I h a v e used t h e latter only casually as a n aid to c o m p o s i t i o n

T h e majority of insects a n d o t h e r types illustrated in t h e line d r a w i n g s a r e placed

in stylized n a t u r a l settings a n d a r e d e ­picted as living animals I have n o t d r a w n figures to scale, except that a n a t t e m p t was

m a d e to indicate c o m p a r a t i v e size, that is,

l a r g e r species a r e l a r g e r t h a n smaller, al­

t h o u g h not in p r o p o r t i o n T h e n e e d occa­

sionally arose to greatly magnify m i c r o ­scopic forms With t h e e x c e p t i o n of t h o s e with t h e n a m e s of o t h e r p h o t o g r a p h e r s o r sources n o t e d in t h e c a p t i o n s , all t h e color

l e n g t h , i.e., front of h e a d to tip of a b d o ­

m e n ) ; B W L (length of insect f r o m front of

h e a d to tips of wings w h e n folded in

r e p o s e ) ; WS (wingspan)

J GENERAL ENTOMOLOGY

E n t o m o l o g y ( G r e e k : entomon insect + logos

discourse) is t h e scientific study of insects A

-1977, 1983) as well as c o m p r e h e n s i v e treat­

m e n t s of a n a t o m y , physiology, classifica­

tion, phylogeny, d e v e l o p m e n t , a n d behav­

ior Because these w o r k s a r e t h o r o u g h a n d widely applicable, it would b e r e d u n d a n t and inefficient to r e p e a t their c o n t e n t s in a specialized b o o k such as this T h e r e m a i n ­

d e r of this c h a p t e r offers a synopsis of

f u n d a m e n t a l topics a n d a n e x p l a n a t i o n of terminology to m a k e t h e book m o r e usable

a n d instructive for g e n e r a l r e a d e r s

References

BARTH, R 1972 Entomología geral Fund

Insto Oswaldo Cruz, Rio de Janeiro

BORROR, D J., AND D M DELONG 1969

Introducáo ao estudo dos insetos Ed Edgard Bluecher, Sao Paulo Brazilian edition trans­

lated and edited by D D Correa et al

BORROR, D J., D M DELONG, AND C H

TRIPLEHORN 1981 An introduction to the

study of insects 5th ed Saunders College, Philadelphia

CARRERA, M 1963 Entomología para vocé Ed

Univ Sao Paulo, Sao Paulo

CORONADO, P., R A MÁRQUEZ, AND A MÁR­

QUEZ 1976 Introducción a la entomología

Ed Limusa, Mexico City

DALY, H V., J T DOYEN, AND P R EHRLICH

1978 Introduction to insect biology and di­versity McGraw Hill, New York

GRASSÉ, P., ed 1949- Traite de zoologie Vols 6-10 Masson, Paris

HAYWARD, K J 1971 Guía para el entomólogo principiante 2d ed Univ Nac Tucumán, Insto Miguel Lillo, Misc no 37: 1-159 KÜKENTHAL, W 1923- Handbuch der zoologie Vol 4 Arthropoda de Gruyter, Berlin LARA, F 1977 Principios de entomología Fac Cien Agr Vet., Univ Estad Paulista "Julio de Mesquita Filho," Sao Paulo

PARKER, S B., ed 1982 Synopsis and classifica­tion of living organisms Vol 2 McGraw-Hill, New York

RICHARDS, O W., AND R G DAVIES 1977

Imms' general textbook of entomology, 10th

ed Vols 1-2 Chapman and Hall, London

RICHARDS, O W., AND R G DAVIES 1983

Tradado de entomología Imms Vol 1, Estruc­tura, fisiología y desarrollo; Vol 2, Clasifi­cación y biología Ed Omega, Barcelona

HISTORY OF LATIN AMERICAN ENTOMOLOGY

Studies o n Latin A m e r i c a n insects a n d related a r t h r o p o d s b e g a n late in t h e his­tory of biology because of t h e belated discovery of t h e New World by E u r o p e a n s

a n d its academic isolation for almost two

c e n t u r i e s thereafter T h e history of e n t o ­mology in t h e r e g i o n is best traced as a series of o v e r l a p p i n g a c c o m p l i s h m e n t s by different categories of s e a r c h e r s a n d for­mats o f investigation, r a t h e r t h a n as t r a d i ­tional chronological p e r i o d s that a r e h e r e not readily identifiable T h e s e categories

a r e largely d e t e r m i n e d by t h e k i n d s of

education available or customary at the

time The earliest disciples were broadly

trained in philosophy, theology, or medi­

cine; later, the narrower disciplines of

natural history, biology, and zoology

evolved Not until very late in the nine­

teenth century did courses in entomology

exist and still later full curricula leading to

a degree in the subject

No general discussion of Latin Ameri­

can entomology is available, although

there are some regional treatments: Lizer y

Trelles (1947) recounts the overall scene

from South America, as do Lamas (1981)

from Peru, Willink (1969) from Argentina,

Kevan (1977) from the West Indies,

Barrera (1955) from Mexico, Fernández

(1978) from Venezuela, and Jirón and

Vargas (1986) from Costa Rica See also

Bodenheimer (1929) and Chardon (1949)

for many basic notes and Howard (1930)

for events in the origins and growth of

practical entomology in most parts of Latin

America Gilbert (1977) provides an index

to published biographies of deceased ento­

mologists (In the following sketch, dates

of birth and death of major figures are

given in brackets [] T h e titles and publica­

tion dates of historically important works

are woven into the text; because they are

well known, they are not cited in the

references.)

Antiquity

There are many evidences of pre-Colum­

bian appreciation for insects, arachnids,

and myriapods among the classic civiliza­

tions of Mexico, Central America, and

northwestern South America Most refer­

ences are to species that affected human

health and welfare Surviving Mayan

(Stempell 1908) and other ancient Mexican

murals and codices depict various species

of economic and religious importance, in­

cluding stingless bees, scorpions, and but­

terflies (Teotihuacán) Early Mexican pot­

tery, also from the Teotihuacán period

(A.D 200-800), are adorned with insect designs, and early Mochica pottery from the northern coast of Peru shows human figures clearly engaged in delousing and infested with the chigoe Other representa­

tions of insect forms appear in sculptures, petroglyphs, and textiles from various cul­

tures (Morge 1973, Tozzer and Allen 1910) Ancient languages and myths con­

tain many entomological allusions, espe­

cially to noxious or ubiquitous species, for example, in Náhuatl, Xochiquetzal, butter­

fly flower goddess (Beutelspacher 1976)

Chroniclers

With the arrival of Columbus, the insects

of the New World became known to West­

ern civilization One might speculate that the lights seen on the shores of Hispaniola, that night of October 11, 1492, were not

native camp fires but glowing Pyrophorns

beetles and thus that an insect was the first thing sighted in America: "After the Admi­

ral had spoken he saw the light once or twice and it was like a wax candle rising

and falling" {J First Voyage)

Among the conquistadors and colonists who followed were scribes and reporters appointed by the Spanish crown to chroni­

cle the discoveries and bring the influence

of Western thinking to the new settle­

ments Many of the sixteenth-century tech­

nical reports of the natural wonders of the newfound lands contained references to

insects One of the earliest, the Historia

General y Natural de las Indias, Islas y firme del Mar Océano (first 20 volumes), was

Tierra-written by Gonzalo Oviedo in 1535 It described for the first time such American curiosities as the cucuyo (headlight beetle), chigger, chigoe, cochineal insect, and sting­

less bees Mentions of the same conspicu­

ous species appeared in other, similar treat­

ments of the period, such as José de

Acosta's Historia Natural y Moral de las Indias (1590) and Bernal Diaz's Historia Verdadera

de la Conquista de la Nueva España

(1568-4 GENERAL ENTOMOLOGY

1632) Bernabé Cobo [1572-1659] wrote

about white butterflies (Ascia monuste) that ittacked crops in Lima in his Historia del

Suevo Mundo (1653), the most important

work of the period on the natural history

of Peru

Fray Bernardino de Sahagún [P-1590]

(fig 1.1) completed his Historia General de

las Cosas de Nueva España in 1560, but it was

not published until the early nineteenth century It described many insects, arach­

nids, and myriapods and was later accom­

panied by illustrations originally intended for it, but from which the text was long

separated {Codex Florentino, fig 1.2.) The

work explained how the Aztecs treated black widow spider bites and scorpion stings and made special mention of useful

Figure 1.1 Fray Bernardino de Sahagún,

post-conquest chronicler of insect life in the New

World (Frontispiece from Historia General de la Cosas de Nueva España, Edition Pedro Ro­

bredo, Mexico, 1938)

Figure 1.2 Figure from Codex Florentino The

stinging arthropod is described by Sahagún in the early sixteenth century as a "scorpion," but in the figure, it is more similar to the larva of

Corydalus cornutus (Megaloptera), called the

"water dog" (perro del agua), an insect widely feared as venomous even today in Mexico

insects such as the maguey worm and the cochineal bug (Curran 1937)

Francisco García Hernández 1578] collected natural objects of medical significance in early colonial Mexico, in­ cluding thirty insects and "worms." His manuscripts were published in various illus­ trated, annotated editions after his death,

[1514-the best known being Rerum Medicarum

Novae Hispaniae Thesaurus sev Planlarum Animalium Mineralium Mexicanorum Historia

(published 1648-1651) (d'Ardois

1959-60, Barrera 1981) in which Tractatus

Quar-lus, Historia Insectorum Novae Hispaniae, was

ostensibly the first unified work on Latin American insects

The chroniclers were savants not schooled in biology or in the methods of scientific investigation Consequently, their statements sometimes contained consider­ able errors, these often the result of believ­ ing too literally the accounts of the Indians But the firsthand recording of natural history by courtiers, travelers, explorers,

HISTORY OF LATIN AMERICAN ENTOMOLOGY 5

traders, soldiers, missionaries, historians,

and adventurers continued in subsequent

centuries (e.g., A de Herrera, Historia Gen­

eral de las Indias Occidentalis [1728]) and

remains a tradition even today (e.g.,

Jacques Cousteau's Amazon Journey [1984])

The creatures described tend to be the

same as those described by earlier authors,

although stinging ants, large centipedes,

Mexican wild silkworms, and tarantulas also

appear

Naturalists

Following the chroniclers onto the scene

were the naturalists, distinguished from

the former by possessing some education

in the biological sciences An early example

was George Marcgrave [1610—1644], who,

during the Dutch invasion of Brazil in

1638—1644, traveled widely and studied

insects in that country An important edi­

tion of his works, citing many indigenous

insects, was published in 1648 by one of his

traveling companions, Guilielmus Piso, in

De Indiae Utriusque re Naturali et Medica Libri

XIV

Following her ten-year stay (1690-1701)

in Surinam, where she collected informa­

tion on insect life histories, Madame Maria

Sybilla Merian [1647-1717] (fig 1.3) pro­

duced her famous Metamorphoses Insectorum

Surinamensium (1705), with superbly done

color plates (Erlanger 1976) The work

contained some errors, including a confu­

sion of the headlight beetles (Pyrophorus),

cicadas, and dragon-headed bugs (Fulgora),

that engendered misconceptions of the

lat-ter's ability to luminesce and stridulate

which persist even today (one plate actually

shows a mongrel insect, a cicada, bearing

the head oí Fulgora) (Frontispiece)

Later naturalists, following in this tradi­

tion and notable for significant observa­

tions on Latin American insects, were

Hans Sloane, A Voyage to the Islands Madera,

Barbadoes, Nieves, St Christophers and Ja­

maica (1707, 1725); G Gardner, Travels in

Figure 1.3 Maria Sybilla Merian, famous for

her observations of insect natural history in the Guianas in the seventeenth century (Frontis­

piece from her botanical work, Erucarum sis ., Amsterdam, 1718)

horten-the Interior of Brazil (1849); Thomas Belt, A Naturalist in Nicaragua (1874); Theodore

Roosevelt, Through the Brazilian Wilderness (1914); Konrad Guenther, A Naturalist in

Brazil (1931); R Hingston, A Naturalist in the Guiana Forest (1932); and others

Renaissance Scholars

The first works on Latin American insects

by those fully qualified as scientists were carried out by established European renais­

sance scholars in absentia They received specimens and reports from collectors and correspondents on the scene but never set foot in the new lands themselves Stingless bees were described in Konrad Gesner's

encylopedic Historia Animaliurn (1607) but

were referenced therein to a work by one Andre Thevet, who "amonst other matters [in the New-found World] reporteth that he

did see a company of flies of Honey-bees about a tree : of which trees there were

a great number in a hole that was in a tree, wherein they made Honey and Wax" (Top-

sel 1967)- In De Animalibus lnsectis

(1602-1618), considered to be the world's first book on entomology, Ulisse Aldrovandi 11522-1605] wrote and figured some Latin American insects, including the cucuyo

This insect, by now famous, also appeared

in Thomas Mouffet's Theatrum Insectorum (1634) alongside a rhinoceros beetle (Mega-

soma) Réné de Reaumur figures and de­

scribes in fine detail a dragon-headed bug

(Fulgora) in his Memoir pour servir a I'histoire desinsectes (17M-1742)

New World specimens were incorpo­

rated into the rapidly growing European collections of the time Nehemiah Grew figures many from the cabinets of the

Royal Society in England (Museum regalis

societatis, 1685)

Culminating this phase of historical development were the great taxonomists, Carolus Linnaeus [1707-1778] and J C

Fabricius [1745-1808], who were able to include a large number of species from the American tropics in their landmark

editions of Systema Naturae (1st ed., 1735;

10th ed., 1758) and various Systemas,

respectively

Collectors

Many of these descriptions were based on specimens provided by a new breed of naturalists to the region, the collectors

Some of the first, who made catching trips to South America in the mid- 1700s, were Pehr Loefling [1729-1756], Carl Dahlberg [1721-1781], Daniel Ro­

insect-u n d e r [1726-1793], and Daniel Solander [1733-1782]

As travel to and conditions in the colo­

nies improved, the number of collectors quickly increased, as did their range (La­

mas 1979, 1981; Papavero 1971, 1973)

The majority of these individuals worked

independently, and many paid their ex­ penses through the sale of their collections

to museums and private collectors in the United States and Europe

A famous duo of pioneer collectors was Karl von Martius [1794-1868] and Johann von Spix [1781-1826] (Fittkau 1983) In 1817-1820, they traversed much of eastern and Amazonian Brazil, collecting thou­ sands of insects that were studied subse­ quently by European specialists A later pair were Osbert Salvin [1835-1898] and Fred­ erick Godman [1834-1919] who traveled widely and amassed specimens in Central America and Mexico in the late 1800s Their extensive collections were assembled

in London, systematically worked up by a number of entomologists, and published in

a lengthy series of volumes under the title

Biología Centrali-Americana (1879-1915), a

classic faunal report Some of the liberes of the infamous French penal colonies in French Guiana from the late 1800s to 1938 made a living by supplying foreign markets with butterflies from the local jungles (Le Moult 1955) Today, many collectors, both commercial and voluntary, continue to pro­ vide material to specialists in their own and other countries

Scientific Expeditions

Other collectors and naturalists partici­ pated in or led the organized scientific or biological expeditions that are a very im­ portant part of the growth of entomologi­ cal science in Latin America These were often sponsored by governments or agen­ cies and included multiple investigators, each with specialized assignments, and were much more elaborate than the simple forays of individuals Primary examples are numerous and date from the early eighteenth century

Antonio de Ulloa [1716-1795] was a military man appointed as Spanish crown representative to the French Académie de Science Expedition to South America in

1735—1746 with La Condamine to mea­

sure the length of an arc of the meridian at

the equator His Noticias Americanas (1772)

contained specific mentions of equatorial

insect life, including an account of a locust

plague The monumental expedition of

the times, however, must be that of Baron

Alexander von Humboldt [1769-1859]

and Aimé Bonpland [1773-1858] to ex­

plore northern South America and Mexico

in 1799-1804 Their extensive insect collec­

tions were researched by Latreille in Eu­

rope (Papavero 1971, chap 4)

Other exemplary expeditions that fur­

thered entomology in Latin America were

several sea voyages with frequent land

stops for collecting, such as the expeditions

of the French vessel La Coquille

(1822-1823), the Swedish Engentes (1851-1852),

and the Austrian Novara (1857-1859) Of

special interest also were the Hamburger

Südperu Expedition in 1936 (Titschack

1951-1954) and the Machris Brazilian Ex­

pedition of 1956 (entomologist F Truxal;

Delacour 1957) A modern example is the

report of entomological results of the

1978-79 Danish Expedition to Patagonia

and Tierra del Fuego (Madsen et al 1980)

Later came those expeditions under­

taken by investigators trained in biology or

zoology who emphasized work with insects

and who conducted studies on their own

collections Three categories of investiga­

tors may be recognized: visiting, expatri­

ate, and native

Visiting Biologists and Zoologists

Perhaps the first biologist to produce sig­

nificant entomological results from his

own excursions in Latin America was Clau­

dio Gay [1800-1873] (fig 1.4), an ambi­

tious French traveler who began to work

with Chilean insects as early as 1836 Later,

he published many research papers, his

most important being Historia Física y Po­

lítica de Chile (arthropod portions,

1849-1852)

About the same time, Charles Darwin

Figure 1.4 Claudio Gay, born in France but

first trained biologist to make a major contribution

to Latin American entomology through his work

in Chile (From portrait in Universidad de Chile;

courtesy of José Valencia)

[1809-1882] made his epic global voyage that included major sojourns in South America He was inclined toward entomol­

ogy and gained some insights into his revo­

lutionary theory of natural selection from observations of South American insects In the initial sentences of his introduction to

the Origin of Species (1859), he states, "When

on board H.M.S 'Beagle,' as naturalist, I was much struck with certain facts in the distribution of the organic beings inhabit­

ing South America [which] throw some light on the origin of species." Some of these facts concerned the distribution of insects and insect examples of sexual se­

lection (e.g., the Chilean stag beetle,

Chiasognatkus) much elaborated in his later

Descent of Man (1871)

Also celebrated among itinerant biolo­

gists of this period was Henry Walter Bates [1825-1892] (fig 1.5) He spent eleven years on the Amazon and its tributaries

8 GENERAL ENTOMOLOGY

Figure 1.5 Henry Walter Bates, first entomolo­

gist explorer of the Amazon Valley in the

mid-18005 (Frontispiece from The Naturalist on the River Amazons, John Murray, London, 1892)

(1848-1859) and collected some 14,000 specimens, including 8,000 species new to science For the first five years of his travels, he was accompanied by Alfred Russel Wallace [1823-1913], who was also

an avid collector but who chose to continue his studies in the Malay Archipelago where

he produced his own theory of natural selection paralleling Darwin's Wallace re­

counts his South American experiences in

A Narrative of Travels on the Amazon and Rio Negro (1853); Bates recounts his in a later

parallel work, The Naturalist on the River

Amazons (1892) Bates collected, but he also

observed and analyzed, producing many papers on Neotropical Coleóptera The work that distinguished him as an ento­

mologist was his Contributions to an Insect

Fauna of the Amazon Valley, Lepidoptera: conidae, published in 1862 (Moon 1976)

Heli-The period of the early to mid-1800s was

a time of independence for most of the countries of Latin America and establish­

ment of national universities, museums, and other learned institutions, with depart­

ments paying attention to terrestrial arthro­

pod life forms Many scholars from Europe emigrated to Latin America Biology came

of age, and considerable progress was made

in the academic phases of entomology, pri­

marily insect systematics But agricultural and medical entomology, knowledge of pes­

ticides, and the role of insects as vectors of disease awaited the threshold of the twenti­

eth century

Other visiting biologists of note were William Beebe [1877-1962], who was gifted with an extraordinary ability of expression and published on many aspects

of Neotropical insect biology, for example,

High Jungle (1949) (Berra 1977), and A S

Calvert, who produced works on Costa Rican insects, including the book, written

with his wife, A Year of Costa Rican Natural

History (1917)

Expatriate Biologists

A special group of early biologists who worked on insects were expatriates They were trained in Europe or North America but were drawn to the Neotropics by its exotic and poorly known insect life They brought with them their education from Western schools and did not merely travel

to the New World but spent the rest of their days in their adopted homes Deserv­ ing special mention in this category is Fritz Müller [1822-1897], born in Germany, who settled in Blumenau, Santa Catarina, Brazil, in 1852 He was a correspondent of Darwin and is best known for his discovery

of the type of mimicry named after him

Another outstanding expatriate biologist was German-born Hermann Burmeister [1807-1892] After a sojourn in Brazil (1850), he settled in Argentina and became the director of the natural history museum

in Buenos Aires and published many im­ portant entomological papers A more mod-

HISTORY OF LATIN AMERICAN ENTOMOLOGY 9

ern example is Felix Woytkowski

[1892-1966], who migrated from his native Po­

land to Peru in 1929 T h e r e he collected

more than a thousand insect species new to

science (Woytkowski 1978) Other notable

expatriate entomologists were Hermann

von Ihering [1850-1930], who emigrated

from Germany to Brazil, Emilio Goeldi

[ 1859-1917], Switzerland to Brazil, Adolfo

Lutz [1855-1940], Germany to Brazil, Paul

Biolley [1862-1909], Switzerland to Costa

Rica, and Henri Pittier [1857-1950], Swit­

zerland to Costa Rica

Native Zoologists

The first full-fledged biologist specializing

in insects who was born in Latin America

was Cuban Felipe Poey [1799-1891] (fig

1.6) He left his birthplace, La Habana, to

study in France but returned to produce

works in entomology in his own land,

especially on Lepidoptera (Centuria de

Lepi-Figure 1.6 Felipe Poey, first native-born Latin

American entomologist (From Memorias de la

Sociedad Cubana de Historia Natural, 1, facing

p 8, 1915)

dópleros de Cuba, 1847) Clodomiro Picado

[1887-1944] also left for study in France, completing his doctoral dissertation on Neotropical bromeliad communities He returned to his native Costa Rica to become its most famous biologist T h e Argentinian Arribálzaga brothers, Felix [1854-1894]

and Enrique [1856-1935], were educated

in their homeland where they carried out extensive studies on insect biology and taxonomy, especially on Diptera

The Entomologists

By reason of their generalized training, the biologists and zoologists could not be con­

sidered entomologists in the strict sense

But because of their scientific abilities, interest, and emphasis on investigation and publication, the title could be logically bestowed on them

Full curricula in the discipline of ento­

mology were not offered in universities until the very late nineteenth century, so professionals in the study of insect biology are virtually all twentieth-century prod­

ucts Their numbers now range in the thousands Who they are and the nature of their accomplishments are best appreci­

ated by reference to the modern literature

and bibliographies such as the Zoological

Record, Parts lnsecta, Arachnida, and

Myri-apoda

The amateur entomologist deserves some special recognition here An active cadre of educated and often highly sophis­

ticated individuals exists who find pleasure

in the study of insects Most are collectors, perhaps the majority working with showy insects like butterflies and beetles, but not always merely for the sake of accumulating specimens Many take advantage of finan­

cial security acquired in other enterprises

to pursue serious questions in entomology

They may even find time to carry out investigations for which the professional finds no support and make valuable contri­

butions directly in their own publications

or in collaboration with professionals

They are therefore distinct from the deal­

ers, whose primary aim in collecting is to profit financially from the sale of their catches

Practical Entomology

Mention of diverse pestiferous Latin Amer­

ican insects is common in the earliest chronicles and later works T h e first refer­

ence to control was made by Francisco

Hernández in his Historia de los Insectos de

Nueva España, written in manuscript in

about the mid-sixteenth century and stat­

ing that the Mexicans used a concoction of tobacco that they spread over the walls to kill fleas in a house (Hoeppli 1969:177) It may have been Henry Hawks, a Vera Cruz (Mexico) merchant, who provided one of the earliest clues to the connection between mosquitoes and human disease, when he wrote in 1572, "This towne is inclined to many kinde of diseases, by reason of the great heat, and a certeine gnat or flie which they call a musquito, which biteth both men and women in their sleepe Many there are that die of this annoyance"

(Keevil 1957)

While evolutionary and taxonomic stud­

ies of insects continued following the birth

of scientific entomology and expanded into the early twentieth century, the discovery of arthropod vectors of animal and human disease and the development of chemical control of crop pests fostered increased work in the applied phases of entomology

in Latin America In medical entomology, major strides were made in the battle against yellow fever and malaria because of the newfound knowledge that mosquitoes were the critical link in the spread of these diseases It was the application of entomo­

logical principles by physicians Carlos Finlay [1833-1915] (fig 1.7), Walter Reed [1851-1902], and William Gorgas [1854- 1920] which rid La Habana of yellow fever

in 1901 and made possible the construction

Figure 1.7 Carlos Finlay, whose ideas led to

the mosquito's role in transmission of yellow fever (From a portrait formerly hung in the Regional Office of the Pan American Health Organization, Washington, D.C.; presently owned by Dr J Fermoselle-Bacardi, Coral Gables, Florida Re­produced with owner's permission)

of the Panama Canal shortly thereafter (Le Prince et al 1916, McCullough 1977) In

1909, Carlos Chagas [1879-1934] demon­ strated that a lethal trypanosome parasite

of humans (Trypanosoma cruzi) was transmit­ ted by a kissing bug (Panstrongylus megistus)

Modern knowledge of agricultural ento­ mology (Doreste et al 1981, Howard 1930), the identification and control of crop pests, was primarily imported, work­ ers and technology in Europe and North America largely determining the course of events Although numerous references to pest insects are scattered throughout the literature of the sixteenth, seventeenth, and eighteenth centuries (Kevan 1977), possibly the earliest scientific investigation into what could really be called economic entomology was made in 1801 when a special commission composed of members

commission) was established in A r g e n t i n a ,

t h e first of m a n y similar agencies, with

ogy was Las Epidemias de las Plantas en la

Costa del Perú by M a n u e l García y M e r i n o

(1878)

Parasitoids a n d p r e d a t o r s of several

pests w e r e i n t r o d u c e d into p r o b l e m areas

with v a r y i n g results by t h e 1930s (Myers

1931), a n d several sites b e c a m e t h e scene

of i m p o r t a n t e x p e r i m e n t a l trials in biologi­

cal c o n t r o l ( H a g e n a n d F r a n z 1973) H o p e s

were realized in t h e Brazilian A m a z o n fly

(Melagonislylum mínense, T a c h i n i d a e ) for

c o n t r o l o f s u g a r c a n e m o t h s T h e sterile

male t e c h n i q u e f o r t h e c o n t r o l o f

screw-w o r m screw-was first tested successfully o n t h e

island of C u r a c a o in 1954

Notes o n t h e history o f t h e various

insects of c o m m e r c i a l value in Latin A m e r ­

ica a r e t o b e f o u n d in t h e systematic

p o r t i o n o f this book (see c o c h i n e a l insects,

silk m o t h , stingless bees, h o n e y b e e , etc.)

References

BARRERA, A 1955 Ensayo sobre el desarrollo

histórico de la entomología en México Rev

Soc Mex Entomol 1: 2 3 - 3 8

BARRERA, A 1981 Notas sobre la interpretación

de los artrópodos citados en el tratado cuarto,

Historia de los insectos de Nueva España, de

Francisco Hernández Fol Entomol Mex 49:

27-34

BERRA, T M 1977 William Beebe, an anno­

tated bibliography Archon, Hamden, Conn

BEUTELSPACHER, C R 1976 La diosa quetzal Soc Mex Lepidop Bol Inf 2: 1-3

Xochi-BODENHEIMER, F S ¡929 Materialien zür Geschichte der Entomologie bis Linné 1: 1 -498; 2: 1-486 Junk, Berlin

CHARDON, C E 1949 Los naturalistas en la

América Latina Sec Agrie, Pec Col., Rep

Dominicana, Ciudad Trujillo

CURRAN, C H 1937 Insect lore of the Aztecs

Nat Hist 39: 196-203

DARDOIS, G S ed 1959-60 Francisco Hernán­

dez, Obras completas Univ Nac México, Mexico City Vol 1 1960, Vida y obra de Francisco Hernández, España y Nueva España en la época de Felipe II by José Miranda

DELACOUR, J 1957 T h e Machris Brazilian Ex­

pedition: general account Los Angeles Co

Mus Contrib Sci 1: 1-11

DORESTE, E., F FERNÁNDEZ, AND P P PAREDES

1981 Contribución a la historia de la entomología agrícola en Venezuela 5th Cong Venezolano Entomol (Maracay), Mem

Pp 29-50

ERLANGER, L 1976 Maria Sybilla Merian, seventeenth-century entomologist, artist and traveler Ins World Dig 3(1): 12-21

FERNÁNDEZ, F 1978 Contribución a la historia

de la entomología en Venezuela Red Fac

Agron (Maracay) 26: 11-27

FITTKAU, E J 1983 Johann Baptist Ritter von Spix, sein Leben und sein wissenschaftliches Werk Spixiana suppl 9: 1 1-18

GILBERT, P 1977 A compendium of the bio­

graphical literature on deceased entomolo­

gists Brit Mus Nat Hist., London

HAGEN, K S., ANDJ M FRANZ 1973 A history

of biological control, in R F Smith, T E

Mittler, and C N Smith, eds., History of entomology Annual Reviews, Palo Alto Pp

433-476

HOEPPLI, R 1969 Parasitic diseases in Africa and the Western Hemisphere, early documen­

tation and transmission by the slave trade

Acta Trop suppl 10: 1-240

HOWARD, L O 1930 A history of applied entomology Smithsonian Misc Coll 84: 1 -

564 (See Pt VI, South and Central America and the West Indies, 417-462.)

JIRÓN, L F., AND R G VARGAS 1986 La

entomología en Costa Rica: Una reseña his­

tórica Rev "Quipu" de Historia de la Ciencia (México) 3(1): 67-77

12 GENERAL ENTOMOLOGY

KEEVIL, J J 1957 Medicine and the Navy 1200-1900 Vol 1 E and S Livingstone, Edinburgh

KLVAN, D K McE 1977

Mid-eighteenth-century entomology and helminthology in the West Indies: Dr James Grainger Soc

Bibliog Nat Hist J 8: 193-222

LAMAS, G 1979 Otto Michael (1859-1934), el cazador de mariposas del Amazonas Col

Suiza Perú Bol 1979(2): 3 5 - 3 8 LAMAS, G 1981 [1980] Introducción a la histo­

ria de la entomología en el Perú Rev

Peruana Entomol 23: 17-37

LE MOULT, E 1955 Mes chasses aux papülons

Ed Pierre Horay, Paris

LE PRINCE, J A., A J ORENSTEIN, AND L O

HOWARD 1916 Mosquito control in Panama

Putnams, New York

LIZER Y TRELLES, C A 1947 Introducción e

historia d e la entomología Argentino Cien

Nat "Bernardino Rivadavia," Publ Ext Cul

Didac 1 (Curso d e entomología) (Buenos Aires), 1-52 (See p 20ff, Historia de la entomología sudamericana.)

MCCULLOUGH, D 1977 T h e imperturbable Dr

Gorgas In D McCullough, T h e path between

the seas: T h e creation of the Panama Canal 1870-1914 Simon & Schuster, New York

Pp 405-426

MADSEN, H B., E S NIELSEN, AND S ODUM,

eds 1980 T h e Danish scientific expedition to Patagonia and Tierra del Fuego 1978-1979

Geogr Tidssk 80: 1-28

MOON! H P 1976 Henry Walter Bates F.R.S

1825-1892, Explorer, scientist and Darwin­

ian Leicestershire Museums, Leicestershire

MORGE, G 1973 Entomology in the Western

world in antiquity and in medieval times In

R F Smith, T E Mittler, and C N Smith, eds., History of entomology Annual Reviews, Palo Alto Pp 3 7 - 8 0

MYERS, J G 1931 A preliminary report on an investigation into the biological control of West Indian insect pests Empire Mrkt Bd

Publ 42:1-173

PAPAVERO, N 1971, 1973 Essays on the history

of Neotropical dipterology, with special refer­

ence to collectors (1750-1905) 2 vols Mus

Zool Univ Sao Paulo, Sao Paulo

STEMPELL, W 1908 Die Tierbilder der handschriften, Zeit Ethnol 1908: 704-743

Maya-TITSCHACK, E., ed 1951-1954 Beitrage zür Fauna Perus: Nach der Hamburger Südperu-Expedition 1936, anderer Sammlungen, wie auch auf Grund von Literaturangaben Vols

1-4 Fischer, Jena

TOPSEL, E 1967 [1658] T h e history of

four-footed beasts and serpents and insects Vol 2

Da Capo, New York

TOZZER, A M., AND G M ALLEN 1910 Animal

figures in the Maya Codices Harvard Univ., Peabody Mus Pap Amer Archaeol Ethnol 4: 273-372, pis 1-39

WILLINK, A 1969 Contribución a la historia de

la entomología Argentina Univ Nac mán, Fund Insto Miguel Lillo Mise 28: 1-30 WOYTKOWSKI, F 1978 Peru, my unpromised land Smithsonian Inst and Nati Sci Found., Washington, D.C

Tucu-LATIN AMERICAN ENTOMOLOGY TODAY

We h a v e seen h o w t h e f o u n d a t i o n s of

e n t o m o l o g y in Latin A m e r i c a w e r e laid,

t h r o u g h four c e n t u r i e s of effort by m a n y types of investigators: c h r o n i c l e r s , g e n e r a l observers, renaissance scholars, collectors,

a n d t r a i n e d entomologists By t h e m i d d l e

of t h e twentieth century, t h e r e was a firm establishment of t h e t r e n d t o w a r d special­ization, b e g u n first with t h e choice of systematists to study c e r t a i n limited taxa, followed by t h e s e p a r a t i o n of t h e a p p l i e d agricultural a n d m e d i c o v e t e r i n a r y fields

a n d m a t u r a t i o n of t h e principles of insect ecology a n d genetics

A n i m p o r t a n t m o d e r n specialization has been t h e s t r o n g interest in tropical biology

by a large n u m b e r of local s t u d e n t s a n d

y o u n g entomologists from N o r t h A m e r i c a

a n d E u r o p e T h e O r g a n i z a t i o n for T r o p i ­cal Studies h a s b e e n a p r i m e m o v e r in this area, principally t h r o u g h t r a i n i n g at field stations in Costa Rica F u n d a m e n t a l discov­eries a r e now b e i n g m a d e a b o u t t h e ecologi­cal a n d evolutionary strategies of insects in the h u m i d lowland e n v i r o n m e n t s , for ex­

a m p l e , t h e t h e o r y of Pleistocene relictual centers of d i s t r i b u t i o n ( B r o w n 1982), t h e theory of island b i o g e o g r a p h y ( M a c A r t h u r

a n d Wilson 1967), a n d t h e o r g a n i z a t i o n of insect societies (Wilson 1971)

T h e vindication of t h e idea of c o n t i n e n ­tal drift (largely from d a t a collected d u r -

LATIN AMERICAN ENTOMOLOGY TODAY 13

ing t h e I n t e r n a t i o n a l Geophysical Year,

B r u n d i n ' s , Transantarctic Relationships and

Their Significance, as Evidenced by

to u n d e r s t a n d t h e diversity of insect life in

Latin A m e r i c a a n d its significance to h u ­

m a n k i n d I n most every country, g o v e r n ­

m e n t s a n d p r i v a t e e n t e r p r i s e a r e recogniz­

ing t h e i m p o r t a n c e of insect f o r m s a n d

e m p l o y i n g entomologists T h e r e h a s b e e n

an increase in t h e n u m b e r s of positions filled by local g r a d u a t e s , a l t h o u g h w o r k e r s

t r a i n e d o r i m p o r t e d f r o m o t h e r c o u n t r i e s still fill m a n y posts E d u c a t e d a m a t e u r s also

r e m a i n i m p o r t a n t c o n t r i b u t o r s Gradually, with t h e h e l p of new technologies for a c q u i r ­ing, r e c o r d i n g , a n d d i s p e n s i n g k n o w l e d g e ,

t h e f a u n a is b e c o m i n g k n o w n , local biologi­

cal p h e n o m e n a a r e b e i n g revealed a n d

i n t e g r a t e d into universal s c h e m e s , losses from d e s t r u c t i o n of food a n d fiber a n d disease a r e b e i n g r e d u c e d , a n d a n a p p r e c i a ­tion for t h e value of insects a n d t h e i r

a r t h r o p o d relatives is b e i n g realized T h e

n e e d r e m a i n s for m o r e facilities a n d fuller staffing of r e s e a r c h institutions a n d g r e a t e r local activity, i n c l u d i n g t h e p o p u l a r i z a t i o n

of insect n a t u r a l history t o t h e g e n e r a l public

References

BROWN, JR., K S 1982 Paleoecology and re­

gional patterns of evolution in Neotropical

forest butterflies In G T Prance, ed., Biologi­

cal diversification in the tropics Columbia Univ Press, New York Pp 255-308

BRUNDIN, L 1967 Insects and the problem of austral disjunctive distribution Ann Rev

Entomol 12: 149-168

HENNIG, W 1950 Grundziige einer Theorie der Phylogenetischen Systematik Deutcher Zentralverlag, Berlin

HILLIS, D M., AND C MORITZ 1990 Molecular

systematics Sinauer Associates, Sunderland, Mass

MACARTHUR, R H., AND E O WILSON 1967

The theory of island biogeography Princeton Univ Press, Princeton

MADSEN, H B., E S NIELSEN, AND S O D U M ,

eds 1980 T h e Danish scientific expedition to Patagonia and Tierra del Fuego 1978—1979

Geogr Tidssk 80: 1-28

SOKAL, R R., AND P H A SNEATH 1963

Principles of numerical taxonomy Freeman, San Francisco

WILSON, E O 1971 T h e insect societies Har­

vard Univ Press, Cambridge

INSECT STRUCTURE AND FUNCTION

T h e physical f o r m a n d b o d y w o r k i n g s (Snodgrass 1 9 3 5 , 1 9 5 2 ; W i g g l e s w o r t h 1984;

C h a p m a n 1982; K e r k u t a n d Gilbert 1984;

King a n d Akai 1 9 8 2 - 1 9 8 4 ; M a n t ó n 1977;

Rockstein 1 9 6 4 - 1 9 7 4 , 1978; Smith 1968;

T r e h e r n e et al 1 9 6 3 - 1 9 8 2 ) of insects a n d their t e r r e s t r i a l a r t h r o p o d relatives a r e as

r e m a r k a b l e a n d c o m p l e x as those of a n y animal t y p e N u m e r o u s s t r u c t u r a l a n d func­

tional systems will b e u s e d in t h e text following as o r g a n i z a t i o n a l topics for a basic review

References

CHAPMAN, R F 1982 T h e insects: structure and function 3d ed American Elsevier, New York

KERKUT, G A., AND L I GILBERT 1984 Com­

prehensive insect physiology, biochemistry and pharmacology Vols 1 — 13 Pergamon, Elmsford, N.Y

KING, R C , AND H AKAI, eds 1982-1984

Insect ultrastructure Vols 1—2 Plenum, New York

MANTÓN, S M 1977 T h e Arthropoda, habits, functional morphology, and evolution Cla­

rendon, Oxford

ROCKSTEIN, M., ed 1964-1974 T h e physiology

of insects Vols 1—6 Academic, New York

ROCKSTEIN, M., ed 1978 Biochemistry of in­

sects Academic, New York

SMITH, D S 1968 Insect cells: Their structure and function Oliver and Boyd, Edinburgh

SNODGRASS, R E 1935 Principles of insect morphology McGraw-Hill, New York

SNODGRASS, R E 1952 A textbook of arthro­

pod anatomy Comstock, Ithaca

TREHERNE, J E., M J BERRIDGE, AND V B

WIGGLESWORTH 1963-1982 Advances in in­

sect physiology Vols 1-16 Academic, New York

WIGGLESWORTH, V B 1984 Insect physiology

8th ed Chapman and Hall, London

Integument

T h e outer, living e p i d e r m i s in insects is a single layer o f generally simple, cuboidal cells t h a t s e c r e t e a n e x t e r n a l n o n l i v i n g cuticle T h e cuticle (Neville 1975) is very

d u r a b l e a n d resistant b e c a u s e of its c o m p o ­sition of w a t e r p r o o f waxes a n d c o m p l e x molecules of such substances as chitin (a

n i t r o g e n o u s polysaccharide) a n d sclerotin (protein) I n c o m b i n a t i o n with t h e e p i d e r ­mis, it forms t h e insect's i n t e g u m e n t ( H e p ­

h a r d areas, o r "sclerites," a r e said to b e well sclerotized Flexibility is allowed by m e m ­

b r a n o u s j o i n t s o r articulations b e t w e e n t h e rigid p o r t i o n s T h u s , t h e i n t e g u m e n t gives

t h e insect its basic f o r m a n d is its p r i m a r y protective system, f o r m i n g a b a r r i e r t o

w a t e r loss a n d e n t r y of p a t h o g e n i c m i c r o o r ­ganisms as well as p r o v i d i n g resistance t o physical t r a u m a

Sclerites m a y also b e s e p a r a t e d by ings, k n o w n as a p o d e m e s (if linear, called

infold-s u t u r e infold-s ; if pitlike, a p o p h y infold-s e infold-s ) It iinfold-s t o t h e

i n t e r n a l p o r t i o n s of a p o d e m e s t h a t t h e

m a i n muscles of m o t i o n a r e a t t a c h e d , giv­ing t h e i n t e g u m e n t a s e c o n d a r y f u n c t i o n ,

t h a t of a skeleton (exoskeleton)

T h e cuticle derives its color n o t only

f r o m its s t r u c t u r a l c o m p o n e n t s b u t f r o m infusions of p i g m e n t s ( C r o w m a r t i e 1959)

a n d m i c r o s t r u c t u r a l d e v e l o p m e n t s (lamel­lae, g r a t i n g s , etc.) t h a t cause s c a t t e r i n g , refraction, a n d d e f r a c t i o n of light waves striking t h e m , r e s u l t i n g in spectral p h e n o m ­

e n a A m o n g t h e p i g m e n t s a r e c o m m o n colored c o m p o u n d s such as m e l a n i n (black), p t e r i n e s (white, r e d , yellow),

c a r o t e n o i d s ( r e d , b r o w n ) , c a r m i n i c acid (carmine), a n d flavones (red, yellow) Physi­cal colors a r e often metallic o r iridescent blues, g r e e n s , a n d r e d d i s h h u e s M a n y

N e o t r o p i c a l butterflies a r e beautifully col­

o r e d f r o m c o m b i n a t i o n s of b o t h pig­

m e n t a r y a n d physical colors localized in t h e wing scales (Ghiradella 1984) Gold a n d silver a r e i n t e r f e r e n c e colors also, b u t u n ­like t h e o t h e r metallics, which a r e p r o d u c e d

by pure, narrow wavelengths, these are

broad-band reflective mixes of radiation

(Neville 1975, 1977) By providing a sur­

face for display of color patterns, the

integument serves additional functions—

protection by crypsis and mimicry, sexual

recognition, and so forth

References

CROWMARTIE, R 1 T 1959 Insect pigments

Ann Rev Entomol 4: 59-76

GHIRADELLA, H 1984 Structure of iridescent

lepidopteran scales: Variations on several

themes Entomol Soc Amer Ann 77:

637-645

HEPBURN, H R., ed 1976 The insect integu­

ment Elsevier, New York

NEVILLE, A C 1975 Biology of the arthropod

cuticle Springer, New York

NEVILLE, A C 1977 Metallic gold and silver

colours in some insect cuticles Ins Physiol

23:1267-1274

Body Cavity

T h e body cavity of all arthropods is not

considered a true coelom, as it lacks a

complete mesodermal lining

Morpholo-gists call it a "mixocoel" because of its

foi ■ ation embryologically from the fusion

of tho blastocoel with parts of the secon­

dary be ' - cavity Because it is filled with

blood, emptying into it from an

open-ended circulatory system, it is also known

as a hemocoel

Segmentation

From their annelid and marine arthropod

ancestors, insects and their terrestrial rela­

tives have inherited a segmented body

Between an anteriormost (acron) and pos­

teriormost segment (telson), a varying

number of segments are interposed, de­

pending on the group T h e r e were origi­

nally 18 (or 19, if a second antennal

segment is recognized) in insects, 19 in

chelicerates, and as many as 100 in

myriapods These were more or less equal

in form and in the possession of a pair of

walking appendages in the first terrestrial

arthropods, much like modern-day centi­

pedes Evolution eventually favored the fusion of adjoining segments (a process called tagmosis) for various functional pur­

poses (e.g., flight in higher insects), and body regions were formed Of these, in­

sects display a triple set, including the head (composed of the acron plus four or five highly fused original segments), a thorax (of three segments, pro-, meso-, and metathorax) and abdomen (with eleven segments, the posteriormost being highly modified into genitalia)

Arachnids and myriapods show differ­

ent patterns of fusion In the former, the head is undefined and its segments totally incorporated into the thorax (cephalo- thorax), which itself may also join into the abdomen, as in mites and ticks Seg­

mentation in these anterior two regions is concealed by a shield or carapace and is evident ventrally only by the serial set of appendages T h e abdomen is either unde­

fined or formed from several segments

Myriapods display only a well-developed head and uniformly segmented thorax- abdomen, with each segment bearing simi­

The Head and Its Appendages

T h e head (Matsuda 1965) is the most highly modified body region, being a sepa­

rate organ (except in arachnids), in which the primitive segmentation is almost oblit­

erated It is normally a rigid capsule, containing the main perceptive and inte- grative neural elements of the animal as well as ingestive organs

T h e many sensory appendages of the

16 GENERAL ENTOMOLOGY

head include the antennae in insects, centi­

pedes, and millipedes, all with one pair

Arachnids lack antennae, their place usu­

ally being assumed by the pedipalps that have become antennalike However, in some arachnids, the pedipalps take other forms and functions, as the claws of scorpi­

ons or walking legs in sun spiders

Around the mouth, modified segmented appendages serve as jaws or stylets for chewing or imbibing liquids and bear food- tasting and smelling organs called palpi In arachnids, these organs are the chelicerae, with the basic scissor form, but they are used directly in feeding by tearing or stab­

bing the food, not chewing T h e chelicerae may lose the movable element and become

a piercing needle in mites, especially para­

sitic ones, and in spiders they are modified into fangs Arachnids use the inner portion

of the leg coxae to scoop liquid nutrient into the simple mouth Among insects and milli­

pedes, there are two pairs of jaws, the anterior mandibles and behind them the maxillae; centipedes have two sets of maxil­

lae In insects, the mandibles and maxillae may retain a primitive toothed or molar form for biting and chewing solid foods, or they may be greatly elongated and bladelike

or hypodermiclike for piercing and siphon­

ing T h e labium may form only a support­

ive sheath around the latter or be itself spongy and absorptive and act directly in food collection Flexibility in adaptation of mouth parts has been a major factor in the success of insects as a group, the variety of morphological types making possible an enormous diversity of food niches and feeding strategies

Although notof appendicular origin, the eyes are of major sensory importance to the head capsule (Horridge 1975) There is a pair of larger multifaceted compound eyes

in adult insects laterally and usually one to three smaller, single-faceted simple eyes medially on top of the head In other terrestrial arthropod groups and immature insects, only simple eyes (ocelli) are present,

either in lateral clumps on the sides of the head (millipedes) or on the back of the cephalothorax (arachnids) Eyes are also often absent altogether (many centipedes)

T h e structure and function of the com­ pound eyes are complex They bulge out

on either side of the head to give a wide range of vision in all directions Each is an aggregation of similar rod-shaped facets called ommatidia, the number of which varies from one per eye in some ants to over 10,000 in dragonflies Each ommatid- ium is composed of elongate sensory cells containing light-sensitive pigments, these concentrated toward the center (thus seen

as a dark rod, called the rhabdome) and exposed on the exterior through a cap­ ping, duplex lens that gathers and focuses light There are also cells with diffuse pigment around the lens T h e sensory cells are nerve cells and are connected directly

to the brain, there being no optic nerve in insects

There are many variations in the de­ tailed structure of the ommatidium, such

as the "apposition" versus the "superposi­ tion" types In the former, the rhabdome is long, and the diffuse pigment cells isolate each ommatidium In the latter, the rhabdome is short, and the screening pig­ ment moves depending on the amount of light in the environment Image formation

is believed to be basically different in the two types, but little is certain about this aspect of eye function It is known that insects generally have good visual acuity and light level accommodation Wave­ length discrimination varies considerably, with a tendency toward the ultraviolet in many species (Silberglied 1979) Many in­ sects, such as bees and butterflies, have good color vision and can orient by polar­ ized light

References

HORRIDGE, G A 1975 The compound eye and vision of insects Clarendon, Oxford

MATSUDA, R 1965 Morphology and evolution

INSECT STRUCTURE AND FUNCTION 17

of the insect head Amer Entomol Inst.,

Mem 4: 1-334

SILBERGLIED, R E 1979 Communication in the

ultraviolet Ann Rev Ecol Syst 10: 3 7 3

T h e insect leg itself is m u l t i s e g m e n t e d

a n d typically c o m p o s e d , f r o m base to tip, of

a coxa ("hip"), t r o c h a n t e r , f e m u r ("thigh"),

tibia ("shin"), a n d t a r s u s ("foot") T h e last

most diverse in t h e insects, a m o n g which

a r e molelike d i g g i n g legs, j u m p i n g legs with

greatly e n l a r g e d , muscle-filled f e m o r a ,

hairy legs for c a r r y i n g p o l l e n , g r a s p i n g a n d

clasping legs in e c t o p a r a s i t e s , a n d flattened,

oarlike s w i m m i n g legs in a q u a t i c insects I n

c e n t i p e d e s , t h e first pair a r e s h a r p fangs

associated with p o i s o n g l a n d s

References

HERREID, C ¥., II, AND C R FOURTNER 1981

Locomotion and energetics in arthropods

Plenum, New York

MATSUDA, R 1970 Morphology and evolution

of the insect thorax Entomol Soc Can., Mem 76: 1-431

WILSON, D M 1966 Insect walking Ann Rev

Entomol I I : 103-122

Wings and Flight

Insects a r e t h e only i n v e r t e b r a t e s with wings T h e s e u n i q u e s t r u c t u r e s a r e n o r ­mally p r e s e n t only in t h e a d u l t stage (may­

flies with w i n g e d s u b i m a g o s b e i n g t h e only exception) a n d always arise f r o m t h e m e s o -

a n d m e t a t h o r a c i c s e g m e n t s T h e i r historic origin is still a d e b a t e d q u e s t i o n , t h e r e being evidence of i n d e p e n d e n t d e r i v a t i o n from t h e body wall a n d their serial

t h r o u g h r e t r o g r a d e evolution o n o n e o r

b o t h s e g m e n t s T h e D i p t e r a a r e c h a r a c t e r ­ized by t h e r e p l a c e m e n t of t h e m e t a ­thoracic p a i r by nonflight, sensory o r g a n s ,

n o u s a n d usually t r a n s p a r e n t , a l t h o u g h it may b e p i g m e n t e d o r c o v e r e d by d e n s e coverings of hairs o r scales It is also

t h i c k e n e d in a linear p a t t e r n to form veinlike struts for s t r e n g t h T h e p a t t e r n of

t h e latter is n o t r a n d o m b u t d e t e r m i n e d phylogenetically a n d is relatively c o n s t a n t

a m o n g taxa (Comstock 1918), t h u s p r o v i d ­ing useful criteria for identification a n d study of relationships Each vein r o o t is

n a m e d , a n d its h o m o l o g u e is recognizable

in all insects So a r e t h e b r a n c h e s of s o m e adventitious veins (such as crossveins) a n d cells f o r m e d by closed sets of veins Slightly different v e n a t i o n a l p l a n s a r e r e c o g n i z e d

by various a u t h o r s (see W o o t t o n 1979)

T h e p r i m a r y p u r p o s e of wings, of

c o u r s e , is flight, a l t h o u g h o t h e r e n d s may

be served T h e y m a y be modified as p r o t e c ­tive shields (such as t h e elytra of beetles

a n d h e m e l y t r a of bugs) o r possess color

p a t t e r n s t h a t p r o v i d e p r o t e c t i o n o r r e c o g n i ­tion signals to o t h e r individuals

T h e flight process in insects h a s b e e n studied extensively a n d f o u n d a e r o d y n a m i -cally u n i q u e (Ellington 1984, G o l d s w o r t h y

a n d W h e e l e r 1 9 8 9 , R a i n e y 1976) All m o v e ­

m e n t is i m p a r t e d by muscles located within the t h o r a x , n o t in t h e wing itself, t h r o u g h a kind of f u l c r u m f o r m e d by t h e lateral b o d y wall a n d elastic m e m b r a n e s a n d sclerotic articulations in t h e wing base T h e s h a p e

a n d t i m i n g of t h e s t r o k e is also d e t e r m i n e d

by t h e s e s t r u c t u r e s It r a n g e s f r o m a slow, simple, u p - a n d - d o w n flapping action as in large butterflies, with a wing beat f r e q u e n c y

of only 4 to 5 p e r s e c o n d , to a c o m p l e x rotational o r twisting m o v e m e n t to a n d fro

S o m e insects such as locusts a n d d r a g o n flies, however, a r e c a p a b l e of sustained flight o v e r l o n g distances for m i g r a t i o n a n d dispersal Small insects f o u n d h i g h in t h e

on air c u r r e n t s , by letting o u t l o n g silk

t h r e a d s g r a s p e d by t h e winds (a process called "ballooning")

GOLDSWORTHY, G f., AND C H WHEELER, eds

1989 Insect flight CRC, Boca Raton

HOCKING, B 1953 T h e intrinsic range and speed of flight of insects Royal Entomol Soc London, Trans 104: 223-345, pi I - V I KUKALOVA-PECK, J 1978 Origin and evolution

of insect wings and their relation to metamor­phosis, as documented by the fossil record, f Morph 156: 53-126

KUKALOVA-PECK, J 1983 Origin of the insect wing and wing articulation from the arthro­pod leg Can J Zool 6 1 : 1618-1669

MATSUDA, R 1981 T h e origin of insect wings (Arthropoda: Insecta) Intl J Ins Morph Embryol 10: 387-398

RAINEY, R C , ed 1976 Insect flight Blackwell, Oxford

WOOTTON, R J 1979 Function, homology and terminology in insect wings Syst Entomol 4:

8 1 - 9 3

The Abdomen and Its Appendages

T h e a b d o m e n is generally t h e least m o d i ­fied of t h e t h r e e b o d y r e g i o n s ( M a t s u d a 1976) It retains its primitive h o m o m e r o u s

s e g m e n t a t i o n in insects, a l t h o u g h in many,

t h e n u m b e r of s e g m e n t s is r e d u c e d

t h r o u g h fusion I n s p i d e r s a n d acarids, only traces of s e g m e n t a t i o n r e m a i n , a n d it

is evanescent in o t h e r g r o u p s M y r i a p o d s have a large n u m b e r of e q u a l s e g m e n t s ,

b u t d i p l o p o d s exhibit a fusion of a d j o i n i n g

s e g m e n t pairs to f o r m d u p l e x , s e c o n d a r y

s e g m e n t s ( h e n c e t h e d o u b l e - p a i r e d legs that give t h e g r o u p its n a m e ) Fully devel­

o p e d a b d o m i n a l walking legs persist only

in this o r d e r a n d t h e C h i l o p o d a A p t e r y gote insects also possess v e n t r a l a p p e n d ­ages o n s o m e basal a b d o m i n a l s e g m e n t s , which r e p r e s e n t vestigial legs

-Also f o u n d at t h e base of t h e a b d o m e n

in varied g r o u p s a r e special modifications

Eggs a r e placed by special a p p e n d a g e s

(ovipositors) B e c a u s e m a n y genitalic struc­

t u r e s a r e often directly involved with t h e

nal sensory a p p e n d a g e s (cerci) t h a t o p e r ­

ate like a kind of r e a r set of a n t e n n a e

References

MATSUDA, R 1976 Morphology and evolution

of the insect abdomen: With special reference

to developmental patterns and their bearings upon systematics Pergamon, New York

SCUDDER, G G E 1971 Comparative morphol­

ogy of insect genitalia Ann Rev Entomol

16: 379-406

TUXEN, S L., ed 1970 Taxonomist's glossary of genitalia in insects 2d ed Munksgaard, Copenhagen

Muscular System

Closely tied functionally to t h e e x o s k e l e t o n

is t h e main m u s c u l a r system All muscles attach to t h e i n t e g u m e n t internally a n d

p r o v i d e m o t i o n to t h e a r t h r o p o d b o d y in all its varied actions T h e y n e v e r f o r m a body wall plexus b u t lie in b u n d l e s r u n n i n g

b e t w e e n insertions T h e latter may be

b r o a d o r a t t e n u a t e d , cover extensive a r e a s

o n sclerites, o r fasten to i n v a g i n a t e d e x t e n ­sions of t h e latter, t h e a p o d e m e s T h e latter, w h e n l o n g a n d slender, a r e ten­

r e p r o d u c t i v e system I n insects a n d t h e i r relatives, all muscle tissue is striated,

capable of slightly m o r e r a p i d twitches

Very r a p i d m o v e m e n t s of insects, such as wing beat f r e q u e n c i e s of 200 to 300 p e r second, a r e m a d e possible by vibratory action of elastic p o r t i o n s of t h e cuticle, t h e muscles themselves c o n t r a c t i n g n o m o r e rapidly p e r s t i m u l u s t h a n t h o s e of a b i r d

T h e t r e m e n d o u s p o w e r p e r b o d y weight

a n d size of m a n y insects (such as t h e giant

h o r n e d scarabs, Dynastes) is also an illusion

S t r e n g t h results from t h e e x e r t i o n of s h o r t fibers a r r a n g e d a l o n g t h e e n t i r e l e n g t h of leg j o i n t surfaces so t h a t t h e load is evenly and widely d i s t r i b u t e d Power o u t p u t a n d metabolic r a t e s of insects, however, a r e

m u c h h i g h e r t h a n in v e r t e b r a t e s , t h e result

of a direct a n d c o n t i n u o u s o x y g e n s u p p l y via t h e t r a c h e a l system

Gross a n a t o m y of t h e m u s c u l a t u r e is highly c o m p l e x , a n d t h e r e may be h u n ­

d r e d s of d i s c r e t e muscles in even a small insect A n early a n a t o m i s t d e s c r i b e d o v e r 4,000 in t h e goat m o t h caterpillar, as c o m ­

p a r e d to a m e r e 5 2 9 in h u m a n s T h i s richness of muscles c o m b i n e d with m e ­chanically d i v e r s e articulations p e r m i t s a diverse r e p e r t o i r e of intricate m o v e m e n t s

orly via t h e a n u s T h e r e a r e t h r e e r e g i o n s

of t h e gut, defined by their e m b r y o n i c origins: a f o r e g u t a n d h i n d g u t , b o t h formed by i n v a g i n a t i o n of t h e blastocoel and lined with e p i d e r m i s a n d cuticle, a n d a

m e s o d e r m a l m i d g u t , lacking a cuticle

T h e r e a r e v a r i o u s diverticula d e p e n d i n g

on t h e g r o u p , m o s t c o m m o n l y a c r o p ( t e m p o r a r y s t o r a g e sac l e a d i n g off t h e esophagus) a n d blind gastric ceca arising

from t h e m i d g u t o r s t o m a c h Salivary

g l a n d s e m p t y into t h e m o u t h cavity o r from t h e tip of a proboscis via l o n g d u c t s associated with t h e h y p o p h a r y n x

Most of t h e digestive e n z y m e s a r e p r o ­

d u c e d by t h e cells in t h e walls of t h e

m i d g u t a n d ceca T h e n a t u r e of these

e n z y m e s varies a c c o r d i n g to d i e t a r y a d a p t a ­tions, p r o t e a s e s a n d lipases p r e d o m i n a t i n g

in carnivores, cellulases a n d related c o m ­

p o u n d s in wood f e e d e r s , k e r a t i n a s e a n d collagenase in scavengers of v e r t e b r a t e connective tissues a n d hair, a n d so o n

Nutrition and Metabolism

Food e n t e r s t h e g u t by t h e m o u t h located o n

t h e front o r u n d e r s i d e of t h e h e a d in insects

a n d m y r i a p o d s o r t h e h e a d region in a r a c h ­nids T h e r e it is m i x e d with p r e d i g e s t i v e

e n z y m e s from t h e salivary g l a n d s , fangs, o r

r e g u r g i t a t i o n s Most digestive processes a r e

r e s e r v e d for t h e i n t e r i o r of t h e s t o m a c h a n d intestine b u t in s p i d e r s begin p r i o r to swal­lowing T h e latter r e g u r g i t a t e o n t h e i r prey,

c a u s i n g enzymatic liquification externally

S o m e early c h a n g e s in food in o t h e r types

of a r t h r o p o d s may b e w r o u g h t by secre­tions of t h e salivary g l a n d s

O n its way to t h e s t o m a c h via t h e e s o p h a ­gus, food may be d i v e r t e d into a c r o p for

s t o r a g e o r g r o u n d u p by a r e g i o n of t h e gut set with spines o r teeth m o v e d by e x t r a heavy muscles ( p r o v e n t r i c u l u s )

T h e n u t r i t i o n a l r e q u i r e m e n t s of insects,

a r a c h n i d s , a n d so o n , a n d t h e i r metabolic processes ( G i l m o u r 1961) also vary e n o r ­mously ( D a d d 1973) T h e s a m e essential

j o r e n e r g y s o u r c e , a n d a l t h o u g h they a r e

INSECT STRUCTURE AND FUNCTION 21

often p r e s e n t in t h e diet, they a r e n o t

always essential a n d c a n b e c o n v e r t e d from

p r o t e i n o r fats T h e r e a r e c o n s i d e r a b l e

differences in t h e ability of d i f f e r e n t in­

sects to utilize p o l y s a c c h a r i d e s Wood

DADD, R H 1973 Insect nutrition: Current

developments and metabolic implications

Ann Rev Entomol 18: 381-420

DOWNER, R G H., ed 1981 Energy metabo­

lism in insects Plenum, New York

GILMOUR, D 1961 T h e metabolism of insects

Freeman, San Francisco

Growth

Food serves n o t only t o p r o v i d e e n e r g y for

activity b u t also to b u i l d u p stores f o r l o n g

d o r m a n t p e r i o d s , a n d , o f c o u r s e , it is t h e

basis f o r g r o w t h As a r t h r o p o d s , with a

confining, a l m o s t n o n e x p a n d a b l e , nonliv­

ing e x t e r i o r cuticle, insects a n d t h e i r kin

achieve size increases a n d m a t u r i t y only by

p e r i o d i c s p u r t s of g r o w t h following molt­

ing T h i s process t a k e s place f r o m a few to

m a n y times d u r i n g t h e a n i m a l ' s life, al­

t h o u g h it ceases after a d u l t h o o d in insects

Molting (or ecdysis) is p r e c e d e d by a cessa­

tion of activity a n d catalysis of t h e lower

cuticle layers w h e n muscles a n d sense o r ­

gans a s s u m e n e w a t t a c h m e n t s O n l y t h e old o u t e r cuticular layer is s h e d , i n c l u d i n g

p e r i o d s of sleep a l t e r n a t i n g with active

l o c o m o t i o n o r f e e d i n g a n d p e r i o d s of sing­

i n g o r c o u r t i n g Events in l o n g - t e r m life cycles a r e also cyclic a n d h a v e i n t e r n a l controls i n t e r a c t i n g with c h a n g e s in a m b i ­

e n t stimuli, day l e n g t h b e i n g a very s t r o n g

o n e (Beck 1980) A l t h o u g h t h e physiologi­

cal basis for such functions is still n o t

u n d e r s t o o d , a n u n d e r l y i n g "biological clock" m e c h a n i s m is p o s t u l a t e d ( S a u n d e r s 1982)

L o n g p e r i o d s of q u i e s c e n c e c o m m o n l y occur in insects a n d relatives, often to c a r r y

t h e a n i m a l t h r o u g h a d v e r s e seasons T h i s

is called d i a p a u s e a n d is characteristic of

h i g h l a t i t u d e o r high elevation species in

t h e w i n t e r t i m e ( h i b e r n a t i o n ) o r of d e s e r t species d u r i n g d r y p e r i o d s (aestivation) A t these times, g r o w t h , d e v e l o p m e n t , a n d activity is a t t e n u a t e d Finally, d i a p a u s e is

b r o k e n with t h e r e t u r n of favorable c o n d i ­tions, a n d e m e r g e n c e o c c u r s S o m e t i m e s

l a r g e n u m b e r s m a y r e t u r n to action simul­

taneously, r e s u l t i n g in p o p u l a t i o n e x p l o ­sions Periods of d o r m a n c y a r e less p r o ­

f o u n d in tropical t h a n t e m p e r a t e insects because of m o r e e q u a b l e e n v i r o n m e n t a l

c o n d i t i o n s in t h e lower latitudes

(Denlin-g e r 1986)

References

BECK, S D 1980 Insect photoperiodism 2d ed

Academic, New York

BRADY, J 1974 T h e physiology of inseci circa­

dian rhythms Adv Ins Physiol 10: 1-115

DENLINCER, D L 1986 Dormancy in tropical insects Ann Rev Entomol 31: 239-264

SAUNDERS, D S 1982 Inseci clocks 2d ed

Pergamon, Oxford

Luminescence

A n o t h e r specialized metabolic j o b to which certain b o d y chemicals a r e p u t is bio-luminescence ( H a r v e y 1957) Q u i t e a n u m ­ber of insects, p r i m a r i l y beetles (glow­

worms, fireflies, h e a d l i g h t beetles, r a i l r o a d worms) a n d millipedes, h a v e evolved light-

p r o d u c i n g o r g a n s (McElroy e t al 1974)

T h e m e c h a n i s m of light p r o d u c t i o n is complex (Case a n d S t r a u s e 1978) b u t basi­

cally involves t h e o x i d a t i o n of luciferin in the p r e s e n c e of t h e e n z y m e luciferase

Luciferin is first activated by A T P in t h e presence of m a g n e s i u m , t h e n oxidized to

an excited f o r m (adenyl-oxy-luciferin) that decays to a lower e n e r g y f o r m with t h e liberation of light T h e r e a c t i o n is cool a n d very efficient, s o m e 9 8 p e r c e n t of t h e energy involved b e i n g released as light

References

CASE, J F , AND L G STRAUSE 1978 Neurally

controlled luminescent systems In P J Her­

ring, ed., Bioluminescence in action Aca­

demic, New York Pp 331-366

HARVEY, E N 1957 A history of luminescence, from the earliest times until 1900 Amer Phil

Soc, Philadelphia

MCELROY, W D., H H SELIGER, AND M D E

-LUCA 1974 Insect bioluminescence Physiol

Ins 2 : 4 1 1 - 4 6 0

Blood and Circulation

All t h e a r t h r o p o d s t h a t a r e t h e subject of this book possess a n o p e n circulatory sys­

tem ( J o n e s 1977) T h a t is, t h e blood moves for t h e m o s t p a r t o v e r a n d a r o u n d t h e tissues a n d o r g a n s , b a t h i n g t h e m a n d e x ­

c h a n g i n g m o l e c u l e s with t h e m directly, in a

c o n t i n u o u s b o d y cavity, t h e h e m o c o e l I n insects, t h e r e a r e n o blood vessels save t h e main a o r t a t h a t leads anteriorly, directly

from t h e h e a r t ( M c C a n n 1970), a n d e m p ­ties into sinuses s u r r o u n d i n g t h e b r a i n I n

c e n t i p e d e s , t h e r e a r e s h o r t lateral a r t e r i e s

l e a d i n g t o t h e g u t a n d o t h e r m i n o r vessels

T h e r e is a " p u l m o n a r y a r t e r y " to t h e b o o k lungs in s p i d e r s as well as s e c o n d a r y vessels

to t h e legs, tail, a n d so o n , in o t h e r a r a c h ­nids T h e h e a r t , which lies dorsally in t h e

h e m o c o e l , j u s t b e n e a t h t h e a b d o m i n a l roof, p r o p e l s t h e blood f o r w a r d with peri­staltic c o n t r a c t i o n s After passing t h r o u g h

t h e b o d y cavity, i n c l u d i n g t h e legs, a n t e n ­nae, wings, a n d o t h e r a p p e n d a g e s , a n d often a i d e d by auxiliary, pulsatile o r g a n s at

bin, such as blood w o r m s , Chironomus) b u t

m a i n t e n a n c e of cellular fluids

References

CROSSLEY, A C 1975 T h e cytophysiology of

insect blood Adv Ins Physiol 11: 117-221

FLORKIN, M., AND C JEUNIAUX 1974 Hemo­

lymph composition Physiol Ins 5: 255-307 JONES, J C 1977 T h e circulatory system of insects C T Thomas, Springfield, 111 MCCANN, F V 1970 Physiology of inseci hearts Ann Rev Entomol 15: 173-200

Hormones

A n i m p o r t a n t class of chemicals t r a n s ­

p o r t e d by t h e blood a r e h o r m o n e s (Novak

1975, Sláma et al 1974) T h e r e a r e m a n y

types, a n d t h e y vary in their effects, even

those f r o m a single e n d o c r i n e o r g a n A few

NOVAK, V J A 1975 Insect hormones Chap­

man & Hall, London

SLÁMA, K., M ROMANUK, AND F SORM 1974

Insect hormones and bioanalogues Springer,

New York

Pheromones

M u c h like h o r m o n e s ( s o m e t i m e s called

" e c t o h o r m o n e s " ) , p h e r o m o n e s ( J a c o b s o n

1972) a r e special k i n d s of biologically ac­

tive substances r e l e a s e d by o n e individual

which cause o t h e r individuals of t h e s a m e species to act in a specific way T h e s e substances a r e e x t r e m e l y n u m e r o u s in kind a n d influence a m o n g insects a n d their relatives In fact, e n t o m o l o g i s t s h a v e realized in r e c e n t years t h a t t h e d o m i n a n t

m e a n s of c o m m u n i c a t i o n b e t w e e n these

c r e a t u r e s is via these m e s s e n g e r substances (Shorey 1976), p e r c e i v e d by olfactory sense o r g a n s , especially on t h e a n t e n n a e ,

c o u r t s h i p a n d c o p u l a t o r y behavior T h e

t r a i l - m a r k i n g substances a n d a l a r m c h e m i ­cals of ants a n d bees t h a t foster a g g r e g a ­tion a r e also p h e r o m o n a l , as a r e t h e caste

a n d activity controlling r e g u l a t o r s in social insect colonies

References

JACOBSON, M 1972 Insect pheromones 2d ed

Physiol Ins 3: 229-276

LEWIS, T., ed 1984 Insect communication

Academic, New York

SHOREY, H H 1976 Animal communication by pheromones Academic, New York

Other External Secretions

A l l o m o n e s a r e c o m p o u n d s p r o d u c e d by insects a n d their relatives t h a t elicit a n t a g o ­nistic reactions b e t w e e n individuals (Bell

a n d C a r d e 1984) T h e y benefit t h e s e n d e r only, usually p r o t e c t i n g it by w a r d i n g off an attack by t h e receiver (Blum 1981) T h e pain-giving (not p r e y - s e d u c i n g ) v e n o m s of female aculeate H y m e n o p t e r a , r e p u g n a n t

o d o r s of m a n y t r u e b u g s a n d beetles, a n d emetic b o d y chemicals (cardiac glycocides

a n d t h e like) in a few butterflies a r e of this category Such also is t h e function of can-

t h a r i d i n (Young 1984a, 19846), a t e r p e n o i d

24 GENERAL ENTOMOLOGY

p r o d u c e d by "blister beetles" (Meloidae)

W h e n p r o v o k e d , these beetles e x u d e blood

c o n t a i n i n g this s u b s t a n c e f r o m t h e tarsal a r t i c u l a t i o n s , a n d they a r e strongly avoided by insectivorous v e r t e b r a t e s a n d

tibio-c a r n i v o r o u s insetibio-cts

O t h e r secretions a r e e x t e r n a l b u t cause

no interactive r e s p o n s e in o t h e r o r t h e same species T h e s e a r e utilitarian sub­

stances involved in t h e life processes of t h e

p r o d u c e r E x a m p l e s a r e silk ( D e n n y 1980) for cocoons a n d webs, adhesives to b i n d

eg g s in place, a n d m a t e r i a l s such as wax or

g u m s for b u i l d i n g s t r u c t u r e s V e n o m used

by spiders, c e n t i p e d e s , s c o r p i o n s , a n d oth­

ers to obtain food also b e l o n g in this category R e g a r d l e s s of function, a r t h r o ­pod v e n o m s a r e usually c o m p a r e d f r o m chemical o r p h a r m a c o l o g i c a l s t a n d p o i n t s (Bettini 1978)

References

BELL, W J., AND R T CARDE, eds 1984

Chemical ecology of insects Sinauer, derland, Mass

Sun-BETTINI, S., ed 1978 Arthropod venoms

Springer, Berlin

BLUM, M S 1981 Chemical defenses of arthro­

pods Academic, New York

DENNY, M W 1980 Silks—their properties and functions Soc Exper Biol., Symp 34: 2 4 7 -

272

YOUNG, D K 1984a Cantharidin and insects:

An historical review Great Lakes Entomol

17: 187-194

YOUNG, D K 19846 Field records and observa­

tions of insects associated with cantharidin

Great Lakes Entomol 17: 195-199

Nervous System

In insects, as with o t h e r a n i m a l s , t h e n e r ­vous tissue is c o m p o s e d of n e r v e cells (neurons), which a r e g r o u p e d into linear nerves a n d g a n g l i a r masses to f o r m a central n e r v o u s system ( T r e h e r n e 1974, Miller 1979), an a u t o n o m i c (or s t o m a t o -gastric) system, a n d a p e r i p h e r a l o r sen­

sory n e r v e system T h e first is v e n t r a l , lying in t h e floor of t h e h e m o c o e l , a n d is characterized by a succession of ganglia

i n t e r s p e r s e d a l o n g a p a i r e d , v e n t r a l n e r v e

c o r d T h e n e r v e cell bodies a r e located

p e r i p h e r a l l y in t h e ganglia, t h e c e n t e r of which a r e o c c u p i e d by a c o m p l e x of n e r v e fibers (the n e u r o p i l e ) t h a t c o n n e c t t h e ganglia as t h e n e r v e c o r d

T h e largest a n d most c o m p l e x g a n g l i o n

is t h e a n t e r i o r m o s t It is dorsal, above t h e

p h a r y n x , in t h e h e a d T h i s is t h e b r a i n ( H o w s e 1970), which may actually be c o m ­

p o s e d of two o r m o r e fused p r i m a r y g a n ­glia It is t h e o v e r r i d i n g c e n t e r of n e u r a l

i n t e g r a t i o n to which t h e o t h e r v e n t r a l g a n ­glia a r e ultimately s u b j u g a t e d , a l t h o u g h each of t h e latter m a y h a v e s o m e d e g r e e of

a u t o n o m y A b e h e a d e d insect m a y con­

t i n u e to live a n d exhibit l o c o m o t o r y a n d sensory activity for s o m e time b e f o r e it eventually dies f r o m such injury

T h e major sensory o r g a n s of t h e h e a d ,

t h e eyes, a n t e n n a e , a n d palpi, a r e con­nected by large nerves directly to t h e b r a i n

T h e b r a i n also c o n t a i n s n e u r o s e c r e t o r y cells a n d functions partly as an e n d o c r i n e

o r g a n as e x p l a i n e d above

T h e first ventral ganglion is also located

in t h e h e a d r e g i o n a n d is associated with ingestive processes T h e r e follows a vary­ing n u m b e r of s e g m e n t e d ganglia, p r i m i ­tively, o n e p e r s e g m e n t , u p to eleven in insects, a n d m a n y m o r e in m y r i a p o d s , b u t

t h e n u m b e r is often less, d u e to fusion of

of t h e e n d o c r i n e system

Efferent n e r v e s r u n from t h e c e n t r a l

n e r v o u s system to t h e muscles in all p a r t s

of t h e body Afferent n e r v e s lead f r o m t h e sensory system, mainly t h e i n t e g u m e n t a r y sense o r g a n s , to t h e c e n t r a l n e r v o u s sys­

t e m T h e cell bodies of sensory n e u r o n s

References

HOWSE, P E 1970 Brain structure and behav­

ior in insects Ann Rev Entomol 20: 3 5 9

-379

MILLER, T A 1979 Insect neurophysiological

techniques Springer, New York

TREHERNE, J E 1974 Insect neurobiology

North Holland, Amsterdam

Integumentary Sense Organs

T h e a r t h r o p o d w o u l d b e isolated from its

of a certain subset o f stimuli i m p o r t a n t to

t h e animal's safety a n d o t h e r life processes

T h e most c o m m o n a n d often most a b u n ­

d a n t sensilla a r e hairlike e x t e n s i o n s (setae,

slight vibrations from air c u r r e n t s o r c o m ­

pression waves a r e p e r c e i v e d For e x a m ­

ple, masses of stretch r e c e p t o r s in t h e

m o t h s a n d g r a s s h o p p e r s , fore tibial h e a r ­ing pits of katydids, a n d acoustical win­

dows in t h e cicada t h o r a x Sensilla a r e often s t r u c t u r e d for t h e

r e c e p t i o n of chemicals in air o r liquids

Such c h e m o r e c e p t o r s (Slifer 1970) usually

h a v e t h i n o r p o r o u s walls so t h e m o l e c u l e s may pass t h r o u g h t h e o u t e r p a r t of t h e

o r g a n a n d stimulate i n n e r receptive sur­

faces T h e y m a y b e e x t r e m e l y sensitive

Calculations for t h e sex a t t r a c t a n t of t h e

d o m e s t i c silk m o t h indicate t h a t a single molecule may elicit a r e s p o n s e

Certain sensilla also react t o a m b i e n t

t e m p e r a t u r e c h a n g e s , r a d i a n t h e a t , p r e s ­

s u r e , humidity, a n d surface m o i s t u r e

(Alt-n e r a (Alt-n d Loftus 1985) P e r c e p t i o (Alt-n of related factors internally a r e by direct cellular sensitivity

References

ALTNER, H., AND R LOFTUS 1985

Ultrastruc-ture and function of insect thermo- and hygroreceptors Ann Rev Entomol 30: 273—

o t h e r insects a n d to t h e h u m a n ear S o m e , such as those resulting from t h e vibration of wings in flight, m a y b e a d v e n t i t i o u s a n d

a p p a r e n t l y h a v e no value t o t h e a n i m a l , b u t most h a v e a specific function a n d o r i g i n a t e from u n i q u e , s o m e t i m e s e l a b o r a t e struc-

tures Extraspecific uses usually a r e to star­

tle a n d a r e p r o t e c t i v e (Masters 1979); specific functions i n c l u d e t h e calling a n d

intra-c o u r t s h i p s t i m u l a t i o n s b e t w e e n t h e sexes,

a g g r e g a t i o n , s p r e a d i n g a l a r m , a n d giving the location of o t h e r colony m e m b e r s in social a n d semisocial f o r m s

S o u n d s m a y b e p r o d u c e d as a by­

p r o d u c t of s o m e activity such as f e e d i n g o r wing m o v e m e n t , t a p p i n g t h e s u b s t r a t e ,

a n d ejections of air, b u t t h e major a n d most effective m e a n s o f sonification involve fric-tional m e c h a n i s m s a n d vibrating m e m ­branes (tymbals) T h e f o r m e r , called stridu-lation, involves two facing surfaces t h a t a r e

r o u g h e n e d a n d that, w h e n m o v e d against each other, p r o d u c e a s o u n d S u c h a r e t h e

n a r r o w s c r a p e r a n d file in t h e base o f t h e fore wings of crickets a n d k a t y d i d s Many

o t h e r insects, beetles, l e p i d o p t e r o u s larvae

a n d p u p a e , a n d so o n , h a v e b r o a d c o r r u ­gated o r r i d g e d a r e a s that w h e n r u b b e d together, give a variety of g r i n d i n g , hiss­

ing, s q u e a k i n g , a n d clicking s o u n d s

S o u n d s p r o d u c e d by t h e vibration o f a

m e m b r a n e d r i v e n by muscles a r e c o m m o n

in H o m o p t e r a , H e t e r o p t e r a , a n d s o m e moths b u t a r e best d e v e l o p e d in male cicadas T h i s s o u n d - p r o d u c i n g o r g a n is located in t h e d o r s o l a t e r a l p a r t of t h e first

a b d o m i n a l s e g m e n t S o u n d is m a d e w h e n the tymbal m u s c l e c o n t r a c t s , p u l l i n g it back rapidly Release allows it t o r e t u r n t o t h e starting position s u d d e n l y against t h e air,

a n d t h e r e s u l t i n g vibrations set u p h i g h intensity air waves t h a t m a y s o u n d to t h e

-h u m a n e a r like a d e a f e n i n g screec-h o r harsh s c r e a m

t u b u l e is o n e cell thick, encircling a lu­

m e n T h e s e cells e x t r a c t waste p r o d u c t s of metabolism f r o m t h e b l o o d , n i t r o g e n o u s

b y - p r o d u c t s usually in t h e f o r m o f uric acid b u t also as u r e a a n d a m m o n i a Potas­sium, s o d i u m , a n d o t h e r i n o r g a n i c ions

a r e also e l i m i n a t e d , a l o n g with a q u a n t i t y

of water

T h e m a i n t e n a n c e of c o n s t a n t salt levels, water, osmotic p r e s s u r e in t h e h e m o l y m p h ,

a n d t h r o u g h t h e cuticle, i n c l u d i n g t h a t lining t h e r e s p i r a t o r y system T h e loss is especially intense in species living in arid

e n v i r o n m e n t s W a t e r is g a i n e d p r i m a r i l y in

t h e food b u t also by d r i n k i n g a n d g e n e r a l

a b s o r p t i o n from h u m i d air Special o r g a n s

of conservation a r e also p r e s e n t in associa­tion with t h e h i n d g u t , w h o s e n o r m a l func­tions i n c l u d e r e a b s o r p t i o n o f w a t e r f r o m

t h e feces O n e of these, t h e c r y p t o n e p h r i d

-i u m , -i n c o r p o r a t e s t h e d-istal e n d s of Malp-i­ghian tubules which loop back o n t o o r into

a t h i c k e n e d p o r t i o n of t h e r e c t u m W a t e r is recycled f r o m t h e latter back into t h e tubules a n d r e u s e d ; feces f r o m these in­sects e m e r g e in a very d r y state

A q u a t i c insects h a v e salt a n d water con­

trol p r o b l e m s d i f f e r e n t from b u t n o less

severe t h a n t h o s e faced by terrestrial types

BARTON-BROWNE, L B 1964 Water regulation

in insects Ann Rev Entomol 9: 6 3 - 8 2

STOBBART, R H., AND J SHAW 1974 Salt and

water balance; excretion Physiol Ins 5:

blood plays n o significant role in this

process e x c e p t in very small, i m m a t u r e

f o r m s t h a t live in d a m p c o n d i t i o n s a n d

aquatics with blood-filled gills T h e tra­

c h e a e o p e n to t h e o u t s i d e t h r o u g h s e g m e n

-tally a r r a n g e d p o r e s , t h e spiracles, which

generally h a v e a closing device to k e e p

water loss to a m i n i m u m L a r g e tubes r u n

S p i d e r s have a t r a c h e a l system in t h e

a b d o m e n only, i n c l u d i n g a variety of m o d i ­fications, a m o n g t h e m "sieve t r a c h e a , "

which a r e l a r g e t r u n k s from t h e e n d s of which o r i g i n a t e n u m e r o u s individual fine

t r a c h e a e Many also possess u n i q u e r e s p i r a ­tory s t r u c t u r e s called "book l u n g s , " which

a r e lamellate, trachealike plates e x t e n d i n g into t h e body cavity Blood flows b e t w e e n

t h e plates, e x c h a n g i n g molecules with t h e

r o u n d i n g liquid, o r c o m b i n a t i o n s of b o t h

A m o n g t h e former, m o s t a r e often associ­

ated a i r stores of o n e k i n d o r a n o t h e r T h e tracheal system itself m a y h a v e sacs o r

e n l a r g e m e n t s to a c c o m m o d a t e air s u p ­plies, o r bubbles m a y b e c a r r i e d b e n e a t h

t h e wings o r held o n t o t h e g e n e r a l b o d y surface by hairs o r o t h e r e x t e n s i o n s of t h e

i n t e g u m e n t P r e v e n t e d f r o m collapse by these extensions, these a i r b u b b l e s act as

"physical gills," o x y g e n a n d c a r b o n d i o x i d e passing in a n d o u t of t h e m t h r o u g h their surface, which acts like a m e m b r a n e ("plastron r e s p i r a t i o n " ) Spiracles c o m m u ­nicating with t h e bubbles t a p t h e air s t o r e

a n d can also function n o r m a l l y s h o u l d t h e

w a t e r d r y u p o r t h e a n i m a l e m e r g e to

a s s u m e a terrestrial p h a s e of existence

Species utilizing a t m o s p h e r i c a i r m u s t

c o m e to t h e surface f r o m time to time t o

r e s t o r e their gaseous provisions, a l t h o u g h

s o m e , such as certain m o s q u i t o larvae, may stay below for very l o n g p e r i o d s of t i m e ,

28 GENERAL ENTOMOLOGY

t a p p i n g air c a r r i e d in t h e vessels of aquatic plants

Small a q u a t i c insects m a y e m p l o y t h e general cuticle as a gill L a r g e types have

o t h e r forms of gill s t r u c t u r e s , e x p a n s i v e nlates o r fingerlike e x t e n s i o n s filled with blood, o r a rich t r a c h e a l n e t w o r k to carry

MILLER, P L 1974 Respiration: Aquatic insects

2d ed Physiol Ins 6: 403-467

Reproduction

Insects a n d like a r t h r o p o d s a r e n o r m a l l y bisexual a n d r e q u i r e sexual c o m m u n i o n o r mating ( B l u m a n d B l u m 1979, T h o r n h i l l and Alcock 1983), with s u b s e q u e n t g a m e t e fusion, for r e p r o d u c t i o n (Davey 1 9 6 5 ,

E n g l e m a n n 1970) Only in a few cases h a s

p a r t h e n o g e n e s i s — a n d in still fewer cases,

h e r m a p h r o d i t i s m — e v o l v e d T h e p r o d u c ­tion of n o r m a l y o u n g by unfertilized fe­

males is p a r t of t h e r e g u l a r r e p r o d u c t i v e process in m a n y H o m o p t e r a , a l t e r n a t i n g with t h e sexual p r o c e s s Unfertilized eggs may b e t h e m e a n s of sex d e t e r m i n a t i o n in others, such as t h e h o n e y b e e , which p r o ­duces d r o n e s by this m e t h o d I n t h e cot­

tony c u s h i o n scale (Icerya purchasi), b o t h

male a n d f e m a l e g o n a d s d e v e l o p in t h e female, a n d self-fertilization takes place

T h e g o n a d s a n d t h e i r i m m e d i a t e d u c t s

a r e almost always p a i r e d T h e g e n e r a t i v e

o r g a n m a y b e single o r m u l t i p l e in myriapods, d e r i v e d f r o m m e s o d e r m a l e m ­bryonic tissue T h e g o n o d u c t s j o i n p a i r e d

or single e c t o d e r m a l i n v a g i n a t i o n s t h a t lead to t h e o u t s i d e via t h e g o n o p o r e T h i s may be located e i t h e r terminally as in m o s t insects o r n e a r t h e base of t h e a b d o m e n in

a r a c h n i d s Male insects a n d m y r i a p o d s usually have

a c o m p l e x set o f genitalia s u r r o u n d i n g t h e

g o n o p o r e , a n e x t e n s i o n of which t e r m i ­nates in a n i n t r o m i t t e n t o r g a n o r p e n i s (often called t h e a e d e a g u s ) T h e s e geni­talia, especially t h e claspers of o n e s o r t o r

a n o t h e r , a r e i m p o r t a n t in locking t h e pair securely a n d precisely t o g e t h e r while t h e penis is i n s e r t e d , f o r m i n g a physical con­nection t h a t is n o r m a l l y species specific ( E b e r h a r d 1985) T h e y m a y also play a

p a r t in physical o r chemical s t i m u l a t i o n necessary for successful c o p u l a t i o n (their

i n n e r surfaces often b e a r sensillar patches) ( A l e x a n d e r 1964) T h e g o n o p o r e is u n -

e l a b o r a t e d in s p i d e r s , t h e function of t h e genitalia b e i n g a s s u m e d by t h e p e d i p a l p s

T h e e x t e r n a l female genitalia a r e rela­tively simple c o m p a r e d to t h e male's, b u t

BLUM, M S., AND N A BLUM 1979 Sexual

selection and reproductive competition in insects Academic, New York

DAVEY, K G 1965 Reproduction in the insects Freeman, San Francisco

EBERHARD, W G 1985 Sexual selection and animal genitalia Harvard Univ Press, Cam­bridge

ENGLEMANN, F 1970 T h e physiology of insect reproduction Pergamon, Oxford

THORNHILL, R., AND J ALCOCK 1983 T h e

evolution of insect mating systems Harvard Univ Press, Cambridge

Fertilization

T h e s p e r m cells p r o d u c e d by t h e testes a r e

i n t r o d u c e d internally into t h e f e m a l e in most forms, t h a t is, fertilization is i n t e r n a l

T h e y m a y b e first k e p t in s t o r a g e in diverticulae of t h e c o m m o n oviduct, how­ever, a n d released to fuse with t h e eggs only as they pass, t h e female t h u s c o n t r o l ­ling t h e time of fertilization

I n t r o d u c t i o n of s p e r m is n o t always directly via t h e g o n o p o r e S e c o n d a r y geni­talia a r e d e v e l o p e d m o s t notably in O d o -

INSECT STRUCTURE AND FUNCTION 29

nata and spiders T h e former transfer the

sperm from the gonopore to the accessory

copulatory organs on the venter of the

third abdominal segment; male spiders use

syringes in the bulbous apex of the

pedi-palps for this purpose Sperm is carried in

a liquid medium, or more commonly, com­

pressed into packets (spermatophores) that

may be inserted into, or formed, in the

common oviduct or its outpocketings

(spermathecae), or are placed on the sub­

stratum to be picked up by the female

Size

Terrestrial arthropods are subject to size

limitations because of the combined restric­

tions of rigidity, lack of permeability, and

weight of the cuticle, which becomes too

much of an encumbrance to movement in

very large forms Also, the diffusion rates of

respiratory gases is insufficient to traverse

the distances necessary through prolonged

tracheal systems, although this is overcome

to some extent by breathing movements

Environmental determinants, such as mois­

ture and food availability, are also impor­

tant (Schoener and Janzen 1968)

In spite of these restrictions, some ex­

tremely large insects are found in Latin

America, all long lived, herbivorous, forest

types In terms of bulk, the record must be

adult males of the large horned scarab, like

Megasoma elephas, which may weigh 40

grams or more Wing expanse is another

measure of size and finds its greatest

expression in the birdwing moth (Thysania

agrippina), with a spread from wing tip to

wing tip of up to 30 centimeters Those

with the longest, although slender, bodies

are the Neotropical centipede Scolopendra

gigantea, which extends 27 centimeters,

and walkingsticks, some 26 centimeters

{Phi Iba losorna phyllinum) from the head to

the tip of the abdomen Indeed, the wet

forests of the Neotropics are traditionally

thought to harbor many insect goliaths

While not the largest overall, some that are

the biggest of their category or impressive

in any sense are many horned beetles such

as Dynastes hércules (17 cm, including horn), morpho butterflies, Morpho hecuba (wing- span 18 cm), tarantulas, Theraphosa lablondi

(20 cm leg span), and lubber grasshoppers,

Tropidacris (wingspan 25 cm, length to

folded wing tips, 13 cm) T h e largest flies

in the world are the Neotropical

Pantoph-thalmus (Pantophthalmidae) that measure 4

centimeters in length and weigh over 2.5 grams

At the low end of the size scale are the smallest known insects, parasitic wasps of

the genus Alaptus (Myrmaridae) with body

lengths of only 0.2 millimeters

Insects and their terrestrial relatives, by and large, are small, the vast majority 6 to

10 millimeters long and 25 to 50 milli­

grams in weight This is their single most important structural characteristic, en­

abling the exploitation of the infinite num­

ber of small niches of nature Insects need little space and minimal sustenance to live and hide from predators

Reference

SCHOENER, T W., AND D H JANZEN 1968

Notes on environmental determinants of tropical versus temperate insect size patterns

Amer Nat 102: 207-224

Genetics and Cytology

Insect genetics has been a fruitful field and has contributed a great deal to this field of general science, particularly through stud­

ies on Drosophila Much of this success is

attributable to the ease with which many insects are maintained in the laboratory, their rapid turnover of generations, diver­

sity of phenotypic expressions of gene effects, and in many cases, giant, well- marked chromosomes

T h e genetic control of a large number

of particular insect characteristics has been elucidated, such as the distribution of dif­

ferent types of hairs, color patterns,

resis-tance to insecticides, and wing venation

Gross changes in Lepidoptera wing color patterns are known to be determined by simple gene differences (Robinson 1971)

Sex in insects is basically determined by (he production of different gametes, al­

though epigenetic factors, such as hor­

mones, are also important (Langé 1970)

Sex chromosomes may be involved, a vari­

ety of combinations being found Males heterozygous XY and XO and females homozygous XX is the usual situation T h e reverse is true of Lepidoptera and Trichop- tera In Hymenoptera, fertilized eggs de­

velop into females, unfertilized eggs into males, the latter therefore being haploid individuals

Genotype and gene frequencies are properties of populations rather than of individual insects Their behavior is impor­

tant to the understanding of evolutionary processes when it is realized that it is shifts

in their frequency, either randomly (ge­

netic drift), by mutation, selection, or exter­

nal events, that lead to speciation and higher order phylogenetic changes A clas­

sic case of the latter is the increase to normalcy of melanism in populations of European moths living in industrial envi­

ronments where heavy soot pollution dark­

ens their resting substrates (Kettlewell 1973) No melanics of this type are yet known in Latin America

Mutations are easily induced in insects

by means of radiation and chemicals T h e former is even used routinely to create sterile individuals for mass release in ge­

netic control schemes (Pal and Whitten 1974)

The mode of gene operation is also becoming known in insects In the giant chromosomes of fly larvae, characteristic swellings, forming after natural hormones contact the cell, appear to indicate activity

of specific genes

Genetic work with other terrestrial ar­

thropod groups aside from insects has lagged behind work with insects

References

KETTLEWELL, H B D 1973 The evolution of

melanism Clarendon, Oxford

LANCÉ, G 1970 Relations entre le nisme génétique du sexe el la controle hor­

détermi-monal de sa differentiation chez les podes: Comparaison avec les vertebres Ann Biol 9: 189-230

arthro-PAL, R., AND M J WHITTEN 1974 The use of

genetics in insect control Holland, Amsterdam

Elsevier/North-ROBINSON, R 1971 Lepidoptera genetics gamon, Oxford

Per-INSECT BEHAVIOR

Insect behavior (Matthews and Matthews 1978) is a rapidly developing field of study that attempts to explain both the complex anatomical and physiological bases and higher, integrative mechanisms for activity Only short-term, decisively determined ac­ tions are recognized in this framework Long-lasting, slowly induced actions, such

as diapause or maturation, are considered physiologic or developmental phenomena (see other parts of this chapter)

Physiochemically and anatomically, in­ sects possess the same elements that con­ trol behavior in all animals Foremost of these is the nervous system (Roeder 1963), including its sensory component, but the muscular and hormonal components play

an essential, if secondary, part It is the degree of complexity of the first that determines the levels on which lines of action lie

A key element of the nervous system in determining behavior is the associative (ad­ juster, internuncial) neuron, which inter­ cedes between receptor (efferent) and effec­ tor (afferent) neurons and has the capacity

to redirect and modify otherwise simple reflex reactions Large numbers of these form masses (neuropiles) in the brain and ventral ganglia and serve as centers of neural integration These are something like the cortex or gray matter of the human brain and define the overall function of a

ganglion They represent the main areas

where activities are generated and orga­

nized A major such center is the corpus

pe-dunculatum ("mushroom-shaped body"),

believed to be the site of summation of

simultaneous excitation from all sources It

tends to be small in arthropods with simple

behavior, large in those with complicated

lives, such as the social Hymenoptera

These cells both stimulate and inhibit

Endocrine secretions are not only caused

to flow in response to nervous command

but are actually part of the nervous system

in the form of neurosecretory cells These

cells produce hormones that move along

the axons and direct other nerve and endo­

crine tissues to emote

Of course, activity is finally the result of

muscular contraction Insects and their rela­

tives may have very large numbers of dis­

crete muscle bundles that predicate a like­

wise elaborate system of efferent nerves It

is fortunate that a lack of obstructive connec­

tive tissue in these animals makes it possible

to dissect and experiment to determine

pathways relatively easily T h e largest

nerves lead to the most active locomotor

organs, the wings and legs Other major

efferents control the mouthparts, anten­

nae, cerci, genitalia, and numerous other

muscularized structures

T h e insect behaviorist looks for chains

or pathways of

stimulation-integration-action to explain activities (Browne 1974)

T h e latter can be considered to be com­

posed of bits or units that meld together

into sequences first, then complexes or

systems T h e simplest movements have the

simplest nerve control and fewest muscles

involved T h e most complex systems have

very large numbers of pathways and pro­

cesses and are so complicated that it is

possible to analyze them only in general

An understanding of the way the whole

insect acts requires an extension of the

rudimentary functioning of the neural,

hormonal, and muscular elements This

extension progresses along a scale of in­

creasing complexity, beginning with called automatic or instinctive behavior and terminating with learned activity

so-The simplest instinctive actions are re­

flex arcs, so-called knee-jerk responses, where a part of the body reacts directly to a stimulus without the intercession of an association nerve An example is the retrac­

tion of the tarsus from a hot surface A step

up from this level occurs when the whole body is coordinated but by nonmodifiable reactions Where only a single action is identifiable, such as movement away from

or toward light or touching or shunning other individuals or objects, the behavior is called a taxis or tropism Such behavior may be positive or negative T h e attraction

of moths to artificial light, the catatonic freezing or "death feigning" display many species use to escape harm, and the follow­

ing of odor trails by dung beetles to find food for their young are specific examples

A series of these tropistic elements may

be strung together, one triggering the next

to form a fixed action pattern These may take up a sizable part of the behavioral repertoires of most insects Pupation in giant silk moth larvae offers an appropri­

ate example: changes in photoperiod or some internal stimulus causes them to cease feeding This initiates defecation and

a wandering, searching activity, leading to the discovery of a suitable pupation site

Even if the latter is not found, the larvae will begin to spin silk and form a cocoon of

a specific shape in which it finally settles and pupates This sequence follows the same steps regardless of changes in exter­

nal stimuli (unless acute) and does not vary according to any information learned by the individual

Insects and other terrestrial arthropods are capable of limited learning (Alloway 1972), defined as any relatively permanent change in behavior that results from prac­

tice Such learning is of a low order and often short lived, but it is often essential to the animal's existence At least two types

32 GENERAL ENTOMOLOGY

have been seen, classical Pavlovian condi­

tioning and, much more commonly, instru­

mental conditioning, where reinforcement stimuli direct the performance of the in­

sect T h e latter is a characteristic especially

of social insects, like the honeybee, which can be trained artifically to fly to a colored surface by food offerings Under natural conditions, this ability is important in re­

cruiting foragers and in efficient utiliza­

tion of a flower nectar food source Some forms, such as cockroaches and ants, facili­

tate to mazes T h e vast majority of these arthropods, however, probably are capable

of virtually no learning whatever

The complexity of some behavior in insects, particularly social insects, most es­

pecially ants, whose lives parallel our own

in some ways, has suggested to some the possibility of the existence of intelligence

As possessed by higher vertebrates, includ­

ing ourselves, no such high degree of learning and reasoning can be truly as­

cribed to these creatures All activity, re­

gardless of how cunning and comprehend­

ing it seems, can be explained on the basis

of fixed action sequences, with very limited learning T h e nesting of digger wasps

(Ammophila) is a classic example: the female

wasp first digs a burrow in sandy soil which

it then closes over at the mouth It then leaves to search for prey, captures it, and returns to the location of the burrow To

do this, it has had to learn a few landmarks

by which it navigates Their misplacement, however, may lead the digger wasp to conclude wrongly on the exact location

The nest, when found, is opened and the prey packed within, an egg is laid on it, and the female exits, closes the nest perma­

nently, and leaves to repeat the process elsewhere All of these are innate, unmodi- fiable acts

The remarkable thing about insect be­

havior is that it may be highly complicated, comparable in this respect alone to verte­

brates, yet it is nearly all controlled by instinctive mechanisms Fundamental life

processes are thus served efficiently, al­ though automatically and unswervingly, and have contributed to their success as a group

It is useful to segregate and classify the kinds of motivation driving the insect body because it is often found that single action sequences operate within them T h e follow­ ing are only representative, as many exam­ ples fit into the categories given; additional types will appear in the main text of this book

1 Alimentation Finding food and feeding

involve specific movements, often elabo­ rate Mosquitoes respond to visual and odor cues to find warm-blooded hosts and then follow tactile stimuli to select a proper station and find a capillary Inter­ nal pressure from expansion of the stomach causes cessation of feeding and induces flight

2 Survival Its host, discovering a mosquito

in the act of feeding, will attempt to destroy or remove it T h e insect displays flight as a survival act, an extremely common one with winged types Other survival-related behavior is shelter seek­ ing, catalepsis, and biting Most protec­ tive coloration is accompanied by pos­ tures that enhance deception or warning patterns

3 Aggression Both intra- and interspecific

agonistic (fighting) behavior occurs in insects, including male-male competi­ tion for females, as in the horned scar­ abs Bees may grapple for a nectary or over territory and females Raiding for food, such as found in many ants, should not be confused with aggression, although the results are the same T h e vanquished colony is perceived as food, not as a rival faction

4 Sex This essential, overriding drive in

all organisms has led to some of the most incredibly complex and even bi­ zarre activities in all groups of terres­ trial arthropods These are divided into

INSECT BEHAVIOR 33

m a t e finding, c o u r t s h i p , c o p u l a t i o n ,

a n d i n s e m i n a t i o n ( T h o r n h i l l a n d

Al-cock 1983)

5 Brood care P a r e n t a l b e h a v i o r occurs in

relatively few insects a n d o t h e r t e r r e s ­

ted a m o n g m e m b e r s of social insect

colonies S o u n d also ties m a n y nonsocial

types t o g e t h e r

7 Tool using It is an a m a z i n g fact that a

few insects actually use tools—in an

tional flight is a c o n s p i c u o u s manifesta­

tion of this behavior, a n d it is most

c o n s p i c u o u s in larger, active f o r m s such

References

ALLOWAY, T M 1972 Learning and memory in insects Ann Rev Entomol 17: 4 3 - 5 6 BROWNE, L B 1974 Experimental analysis of insect behavior Springer, New York

MATTHEWS, R M., AND J R MATTHEWS 1978

Insect behavior Wiley, New York

ROEDER, K D 1963 Nerve cells and insect behavior Harvard Univ Press, Cambridge

THORNHILL, R., AND J ALCOCK 1983 T h e

evolution of insect mating systems Harvard Univ Press, Cambridge

DEVELOPMENT AND LIFE CYCLES

Eggs

W h e t h e r e x t e r n a l to t h e female p a r e n t ' s

b o d y (oviparity) o r t e m p o r a r i l y within parity), all insects, s p i d e r s , a n d allied t e r r e s ­trial a r t h r o p o d s start their lives as eggs ( H i n t o n 1981) Eggs c o m e in a n a m a z i n g variety of s h a p e s a n d sizes T h e y a r e usually placed singly or in g r o u p s in p r o x i m i t y to

(vivi-t h e juvenile's food s o u r c e b u (vivi-t m a y b e sca(vivi-t­

t e r e d indiscriminately only in t h e g e n e r a l habitat w h e r e d e v e l o p m e n t o c c u r s M a n y have e l a b o r a t e cuticular s c u l p t u r i n g , a n d

s o m e possess devices for a t t a c h m e n t to t h e

s u b s t r a t u m o r caps (opercula) that o p e n to allow egress of t h e y o u n g A n u m b e r of species protect their eggs f r o m m o i s t u r e loss a n d t r a u m a by c o v e r i n g t h e m with froth o r e n c a s i n g t h e m in o t h e r substances

t h a t h a r d e n a r o u n d t h e m (oothecae)

Reference

HINTON, H E.,ed 1981 Biology of insect eggs

Vols 1-3 Pergamon, Oxford

Embryology

fust p r i o r to fertilization, insect eggs a r e

c o m p o s e d mostly of yolk a n d small islands

of cytoplasm s u r r o u n d i n g t h e female n u ­cleus on o n e e d g e W h e n t h e e g g is laid, the nucleus is usually in t h e m e t a p h a s e of the first meiotic division, in which state it receives t h e s p e r m , o n e of which u n i t e s with t h e oocyte after meiosis is c o m p l e t e

T h e nucleus t h e n m i g r a t e s to t h e c e n t e r of the egg a n d begins to divide mitotically

T h e r e s u l t i n g cells m o v e to t h e p e r i p h e r y and form t h e b l a s t o d e r m , o r early e m b r y o , which later l o d g e s o n o n e side of t h e egg

T h e g e r m layers a n d e m b r y o n i c m e m ­branes soon d e v e l o p , a n d d e t e r m i n a t i o n of

s e g m e n t a t i o n a n d t h e p r i m a r y o r g a n s a n d tissues e n s u e s T h e a p p e n d a g e s a p p e a r , and after a t i m e , t h e perfect b o d y of t h e first j u v e n i l e stage is c o m p l e t e T h i s stage takes d i f f e r e n t f o r m s d e p e n d i n g o n t h e evolutionary level of t h e g r o u p Fairly similar e m b r y o l o g i c a l steps a r e followed by

o t h e r terrestrial a r t h r o p o d s ( J o h a n n s e n and Butt 1941) A major e x c e p t i o n a r e t h e springtails (Collembola), w h o s e eggs u n ­

d e r g o holoblastic cleavage

Reference

JOHANNSEN, O A., AND E H BUTT 1941

Embryology of insects and myriapods Mc­

Graw-Hill, New York

g r o u p s , s e g m e n t s a r e a d d e d as d e v e l o p ­ment p r o c e e d s

As t h e a n i m a l p r o g r e s s e s t o w a r d m a t u ­rity, it increases in size, a n d c h a n g e s in

i n t e r n a l a n d e x t e r n a l f o r m a n d p r o p o r ­tions o c c u r to a g r e a t e r o r lesser d e g r e e (Sehnal 1985) In m o s t n o n i n s e c t s a n d primitive a p t e r o u s insects, t h e i m m a t u r e s

a r e fairly similar to t h e a d u l t s J u v e n i l e insects of t h e h i g h e r o r d e r s t h a t possess wings, however, u n d e r g o a fair a m o u n t of

b o d y modification, called m e t a m o r p h o s i s , primarily associated with t h e g r o w t h of t h e wings a n d e x p l o i t a t i o n of habitats differ­

e n t from t h e a d u l t M e t a m o r p h o s i s is said

to be " g r a d u a l " (incomplete) in lower

w i n g e d insects with e x t e r n a l l y d e v e l o p i n g wing b u d s ; t h e single j u v e n i l e type is called

a n y m p h (or s o m e t i m e s n a i a d in aquatics)

N y m p h s generally have f e e d i n g a n d o t h e r habits similar to t h e a d u l t ; n a i a d s live

r a t h e r d i f f e r e n t lives b e c a u s e of t h e i r wa­ter habitats M e t a m o r p h o s i s is " c o m p l e t e "

t h e e v o l u t i o n a r y success of these insects

t h r o u g h t h e d i c h o t o m o u s specialization of life functions ( f e e d i n g a n d g r o w t h by im­

m a t u r e s , dispersal a n d r e p r o d u c t i o n by adults) Divergence of b o d y f o r m a n d func­tion has even t a k e n a f u r t h e r s t e p in m a n y species with v a r y i n g types of larvae ( h y p e r -

m e t a m o r p h o s i s ) such as f o u n d in t h e blis­ter beetles (Meloidae), chalcidoid wasps,

a n d o t h e r s I m m a t u r e s of different insect

g r o u p s a r e called by v a r i o u s n a m e s For

e x a m p l e , l a r v a e of L e p i d o p t e r a a r e cater­pillars; p u p a e of butterflies, chrysalids; larvae of muscoid flies, m a g g o t s ; a n d bee­tle larvae, g r u b s P u p a e generally a r e p r o ­tected by their location, u n d e r g r o u n d in cells o r in wood o r o t h e r m a t e r i a l or

e n c a s e d in a cocoon of silk s p u n by t h e

p r e p u p a l instar

References

AGRELL, I P S., AND A M LUNDQUIST 1973

Physiology and biochemical changes during

insect development Physiol Ins 1: 159-247

SEHNAL, F 1985 Morphology of insect develop­

ment Ann Rev Entomol 30: 89-109

m e n t constitutes its life cycle ( T a u b e r et al

1985) Life cycles a r e as varied as t h e kinds

(bivoltine) o r m u l t i v o l t i n e , with two to sev­

eral g e n e r a t i o n s p e r year T h e latter a r e

m o r e typical of tropical o r o t h e r stable

T h e females lay eggs t h a t h a t c h i n t o t h e asexual forms o n c e again P r o d u c t i o n of sexual f o r m s is c o n t r o l l e d by c h a n g e s in

t e m p e r a t u r e a n d p h o t o p e r i o d ; u n d e r con­

stant tropical c o n d i t i o n s , cyclical a l t e r n a ­tion of g e n e r a t i o n s m a y n o t occur

Reference

TAUBER, M J., C A TAUBER, AND S MASAKI

1985 Seasonal adaptations of insects Oxford Univ Press, New York

EVOLUTION AND CLASSIFICATION

T h e r e c o n s t r u c t i o n of t h e historical evolu­

tion a n d d e t e r m i n a t i o n of t h e i n t e r r e l a t i o n ­ships of t h e p r e s e n t l y e x t a n t o r d e r s of insects a n d o t h e r terrestrial i n v e r t e b r a t e s has n o t b e e n settled by a n y m e a n s T h e r e

36 GENERAL ENTOMOLOGY

r e m a i n m a n y c o n t r o v e r s i e s , e v e n o v e r m a ­jor theses, such as t h e m o n o p h y l y (descent from a single a n c e s t r a l line) of t h e A r t h r o -poda o r of t h e a p t e r y g o t e h e x a p o d s

T h e r e is e x t e n s i v e l i t e r a t u r e o n these dis­

a g r e e m e n t s a n d r e l e v a n t a r g u m e n t a t i o n ( A n d e r s o n 1 9 7 3 ; B o u d r e a u x 1979; G u p t a 1979; M a n t ó n 1977; S h a r o v 1966)

T h e a r t h r o p o d g r o u p s i n c l u d e d in this book a r e all basically t e r r e s t r i a l , p r o b a b l y

by way of several i n d e p e n d e n t , parallel evolutionary p a t h w a y s , from varied p r e c u r ­sors a m o n g t h e O n y c o p h o r a , C r u s t a c e a (Isopoda), U n i r a m i a ( m y r i a p o d s a n d in­

sects), a n d C h e l i c e r a t a ( a r a c h n i d s ) , a n d a r e thus only distantly r e l a t e d ( M a n t ó n 1977:

2 5 7 - 2 5 8 )

T h e o n y c o p h o r a n line s e e m s to attach most closely to t h e m y r i a p o d a n , a n d these animals c a n n o l o n g e r b e c o n s i d e r e d inter­

mediate phylogenetically b e t w e e n a n n e l i d s and a r t h r o p o d s , t h e latter n o w b e i n g recog­

nized as a polyphyletic g r o u p T h e y a r e not ancestral to e i t h e r t h a t g r o u p o r

H e x a p o d a , n o r is t h e latter d e s c e n d e d from t h e M y r i a p o d a Embryological evi­

dence indicates t h a t all t h r e e h a v e di­

T h e chelicerates, d i s t i n g u i s h e d f u n d a ­mentally by t h e i r c h e l i c e r a t e m o u t h p a r t s , are virtually all t e r r e s t r i a l , a l t h o u g h p r o b a ­bly d e r i v e d f r o m originally m a r i n e ances­

tors Evolution within t h e s u b p h y l u m is n o t clear All efforts to s u b d i v i d e t h e o r d e r s have r e m a i n e d inconclusive, as h a v e associ­

ated p h y l o g e n e t i c s p e c u l a t i o n s T h o s e with book l u n g s ( S c o r p i o n i d a , U r o p y g i , Ambly-pygi, spiders) p r e s u m a b l y can b e g r o u p e d ; scorpions, with their c o m p l e t e s e g m e n t a ­tion, a r e t h e most p r i m i t i v e I n b o d y s h a p e and e x t e r n a l genitalia, t h e O p i l i o n e s r e s e m ­

ble s o m e primitive mites, with which t h e y seem to form a close b r a n c h T h e o t h e r

g r o u p s a r e all isolated

T h e p h y l o g e n y of t h e primitively m a n

-d i b u l a t e U n i r a m i a ( a p p e n -d a g e s with sin­gle stem) is fairly well u n d e r s t o o d , at least for t h e insects in g e n e r a l (Kristensen 1981) M y r i a p o d s r e t a i n h o m o m e r i s m ,

h a v i n g only a distinct h e a d , b u t s e e m to possess t h e basic b o d y s t r u c t u r e likeliest to

p r e c e d e that of insects T h e a n c e s t o r s of

t h e insects ( H e n n i g 1981) evolved a t h r e e somite t h o r a x a n d t h r e e p a i r s of legs at a n early time, r e d u c i n g t h e m a n y e q u a l b o d y

o v e r g r o w n by cranial folds ( e n t o g n a t h y )

T h e r e a r e also wingless ( A p t e r y g o t a ) p r e ­decessors in b o d y d e s i g n to t h e d o m i n a n t insects that evolved wings early in t h e i r history (Pterygota) (Kukalová-Peck 1987)

At first (Paleoptera), wings w e r e clumsy,

o u t w a r d l y projecting, fixed, flight o r g a n s ,

as seen in m a n y extinct g r o u p s of t h e Paleozoic (e.g., P a l a e o d i c t y o p t e r a ) a n d ex­

t a n t mayflies a n d O d o n a t a , b u t soon ac­

q u i r e d i m p r o v e m e n t s , a m o n g t h e m t h e ability to be flexed o v e r t h e b o d y which all

t h e h i g h e r o r d e r s h a v e ( N e o p t e r a ) E v e n those that have secondarily lost wings alto­gether, often in association with e c t o p a r a -sitism (fleas, lice, b e d b u g s , etc.), r e t a i n t h e thoracic s t r u c t u r e of their fully w i n g e d ancestors

T h r e e major lines e m e r g e d within t h e

h i g h e r w i n g e d insects T h e first, most p r i m ­itive a s s e m b l a g e ( P o l y n e o p t e r a ) , which

s o m e w o r k e r s q u e s t i o n as m o n o p h y l e t i c , includes t h e " o r t h o p t e r o i d " g r o u p s , t h e

O r t h o p t e r a , G r y l l o p t e r a , D e r m a p t e r a , a n d

o t h e r o r d e r s t h a t display a n e n l a r g e d , fanlike h i n d wing with m a n y l o n g i t u d i n a l veins, m u l t i s e g m e n t e d tarsi, a n d m a n y

M a l p i g h i a n t u b u l e s a n d ganglia internally

EVOLUTION AND CLASSIFICATION 37

(* = groups not covered in this book; included

for reference only.)

O r d e r Thysanura—silverfish

a n d bristletails Infraclass Pterygota—winged insects

S u p e r o r d e r P a l e o p t e r a ancient—winged insects

-O r d e r E p h e m e r o p t e r a — mayflies

O r d e r A n o p l u r a - s u c k i n g lice

O r d e r H e m i p t e r a — t r u e b u g s ( h e t e r o p t e r a n s a n d

References

ANDERSON, D T 1973 Embryology and eny in annelids and arthropods Pergamon, Oxford

phylog-BOUDREAUX, H B 1979 Arthropod phylogeny, with special reference to insects Wiley, New York

GUPTA, A P., ed 1979 Arthropod phylogeny Van Nostrand Reinhold, New York

HENNIG, W 1981 Insect phylogeny Wiley, Chinchester, Eng

KRISTENSEN, N P 1981 Phylogeny of insect orders Ann Rev Entomol 26: 135—157

KUKALOVA-PECK, J 1987 New Carboniferous

Diplura, Monura, and Thysanura, the pod ground plan, and the role of thoracic side lobes in the origin of wings (Insecta) Can J Zool 65: 2327-2345

hexa-MANTÓN, S M 1977 T h e Arthropoda, habits, functional morphology and evolution Claren­don, Oxford

SHAROV, A G 1966 Basic arthropodan stock with special reference to insects Pergamon, Oxford

FOSSIL INSECTS

K n o w n Latin A m e r i c a n fossil insect sites

a r e few, b u t they h a v e p r o d u c e d c o n s i d e r ­able material r e p r e s e n t i n g several types of fossilization Most r e p r e s e n t relatively r e ­cent strata (Cenozoic)

I m p r e s s i o n s in s e d i m e n t a r y rock f r o m

t h e E o c e n e in S o u t h A m e r i c a a r e m o s t significant (Martinez 1982) O n e of t h e best

k n o w n beds is f o u n d at S u n c h a l , A r g e n t i n a ,

in t h e p r o v i n c e of Jujuy M a n y s p e c i m e n s of weevils a n d o t h e r insects w e r e e x c a v a t e d

t h e r e by e n t o m o l o g i s t T D A Cockerell early in this century T h e oldest insects f r o m

t h e region a r e of an u n i d e n t i f i e d o r d e r

(Eugeropteron a n d Geropteron) from m i d d l e

C a r b o n i f e r o u s b e d s in t h e Sierra d e los Llanos of t h e p r o v i n c e of Rioja, also in

A r g e n t i n a O t h e r i m p o r t a n t sites of insect

fossils p r e s e r v e d in s e d i m e n t a r y rocks a r e

located in Rio G r a n d e d o Sul, Brazil, a n d at

Bajo d e Veliz, in t h e p r o v i n c e o f San Luis in

time a n d r e m a i n e d clear within so t h a t t h e

most m i n u t e s t r u c t u r e s (hairs, genitalia,

I n t h e A m e r i c a s , t h e r e a r e also a n u m b e r

of o t h e r k n o w n b u t u n e x p l o r e d a m b e r deposits, for e x a m p l e , in C o l o m b i a (Cocker-ell 1923), Brazil (Froes A b r e u 1937), a n d surely o t h e r c o u n t r i e s (Poinar a n d A g u d e l o 1980)

G o o d p r e s e r v a t i o n is also characteristic

of t h e Q u a t e r n a r y A g e r e m a i n s f o u n d in asphalt deposits S o m e sites in this c a t e g o r y

a r e located in T r i n i d a d (Blair 1927) a n d at Talara o n t h e n o r t h e r n P e r u v i a n coast ( C h u r c h e r 1966) H e r e , b e c a u s e o f t h e stickiness of t h e tarlike m e d i u m a n d t h e attractiveness of t h e surface, which looks like water, asphalt seeps f o r m very efficient small a n i m a l t r a p s T h e most n u m e r o u s kinds of insects f o u n d as fossils in these deposits a r e h a r d - b o d i e d g r o u n d beetles, aquatics, a n d c a r r i o n f e e d e r s Aquatic in­

sects a r e s o m e t i m e s indicative of t h e p r e s ­ence of freshwater pools n e a r t h e a s p h a l t

o r overlying it T h e y w e r e e n t r a p p e d w h e n

t h e water d r i e d u p d u r i n g d r o u g h t peri­

ods C a r r i o n - f e e d i n g species w e r e c a u g h t

a l o n g with t h e carcasses of v e r t e b r a t e s t h a t died in t h e black q u a g m i r e s

Because of t h e i r small size a n d delicate ness, insects a n d t h e i r relatives p r o d u c e

-g o o d fossils only in fine grained o r h o m o ­

g e n e o u s matrices T h e f o r e g o i n g a r e of this type O t h e r m o d e s of fossilization t h a t

m a y b e i m p o r t a n t in Latin A m e r i c a , a n d which have b e e n scarcely investigated, a r e

p e r m i n e r a l i z a t i o n (such as in m i n e r a l

-c h a r g e d waters), p e a t a n d soft -coal e n -c a p ­sulation, cave s e d i m e n t s (Miller 1986), a n d silicification, especially e v i d e n t in calcare­

ous n o d u l e s Evidences of f e e d i n g , b o r i n g ,

40 GENERAL ENTOMOLOGY

coprolites, a n d trails s h o u l d also b e c o m ­

m o n in d e p o s i t s o f p l a n t fossils Insect remains in association with a n c i e n t h u m a n remains m a y also b e of c o n s i d e r a b l e a r ­chaeological significance (e.g., W a r n e r a n d Smith 1968)

References

BARONI-URBANI, C B 1980 First description of fossil gardening ants Amber collection Stutt­

gart and Natural History Museum Basel:

Hymenoptera: Formicidae I: Attini Stutt

Beitr Naturk Ser B (Geol Paleon.) 54: 1-13

BARONI-URBANI, C B., AND J B SAUNDERS

1982 T h e fauna of the Dominican amber:

The present state of knowledge 9th Carib

Geol Conf (Santo Domingo, 1980) Trans 1:

213-223

BLAIR, K G 1927 Insect remains from oil sands in Trinidad Entomol Soc London Trans 75: 137-141

CHURCHER, C S 1966 T h e insect fauna from the Talara tar-seeps, Peru Can J Zool 44:

985-993

COCKERELL,T D A 1923 Insects in amber from South America Amer J Sci 5: 331-333

COKENDOLPHER, J C 1986 (1987) A new species

of fossil Pellobunus from Dominican Republic

amber (Arachnida: Opiliones: dae) Carib J Sci 2 2 : 2 0 5 - 2 1 1

Phalangodi-FROES ABREU, S 1937 Sobre a ocorréncia de

ambur nos arenitos da serie Bahia: Brasil

Inst Nac Tech (Rio de Janeiro) Bol Inf

2(4): 8

GRIMALDI, D A., ed 1990 Insects from the Santana Formation, Lower Cretaceous, of Brazil Amer Mus Nat Hist Bull 195: 1 -

191

HÜNICKEN, M A 1980 A giant fossil spider

(Megarachne servinei) from Bajo de Veliz

Acad Nac Cien Córdoba, Bol 53: 317-325

HURD, JR., P D., R F SMITH, ANDJ W DURHAM

1962 T h e fossiliferous amber of Chiapas, Mexico Ciencia 21(3): 107-118, PI I—II

MARTÍNEZ, S 1982 Catálogo sistemático de los insectos fósiles de América del Sur Fac

Hum Cien (Univ Rep., Montevideo) Ser

Cien Tierra, Rev 1(2): 2 9 - 8 3 MILLER, S E 1986 Phylum Arthropoda, Class

Insecta In D W Steadman, Holocene verte­

brate fossils from Isla Floreana, Galápagos

Smithsonian Contrib Zool 413: 1-103

POINAR, JR., G O., AND F AGUDELO 1980 El

ámbar: Oro fósil del nuevo mundo Americas 32(10): 3 3 - 4 0

POINAR, JR., G O., AND R HESS 1982

Ultra-structure of 40-million-year-old insect tissue Science 215: 1241-1242

RICE, P C 1979 Amber of Santo Domingo— mining in the Dominican Republic Lapidary

J (Nov 1979): 1804-1810

RICE, H E., AND P C RICE 1980 Pepitas de sol

antillano Americas 32 (10): 3 7 - 4 1

SANDERSON, M W., A N D T H FARR 1960 Amber

with insect and plant inclusions from the Dominican Republic Science 131: 1 3 1 3 -

1314

VARIOUS AUTHORS 1963, 1971 Studies of fos­

siliferous amber arthropods of Chiapas, Mex­ico, Pts 1, II Univ Calif Publ Entomol 3 1 : 1-53, pis 1-3; 63: i-vi, 1-106, pis 1-3

WARNER, R E., AND G E SMITH, JR 1968 Boll

weevil found in pre-Columbian cotton from Mexico Science 162(3856): 911-912

WILSON, E O 1985 Invasion and extinction in the West Indian ant fauna: Evidence from the Dominican amber Science 229(4710): 265-267

INSECT NAMES

All k n o w n o r g a n i s m s , i n c l u d i n g insects

a n d their relatives, h a v e a scientific n a m e ,

a n d m a n y also h a v e a c o m m o n n a m e (Goto 1982)

Scientific n a m e s a r e a p p l i e d a c c o r d i n g

to r i g o r o u s p r o c e d u r e s (Ride et al 1985), with consistency, universality, a n d stability

by publication T h e first d e s c r i b e r is e n t i ­tled to a u t h o r s h i p , a n d all o t h e r s a r e obliged to u s e t h a t n a m e T h e t e r m "new species" r e f e r s to o n e t h a t h a s b e e n so

f o u n d for t h e first t i m e , n o t to freshly

INSECT NAMES 41

plied to t h e insects a n d t h e i r relatives in

all Latin A m e r i c a n c o u n t r i e s Léxica have

a c c o r d i n g to n o consistent set of stan­

d a r d s , v a r y i n g f r o m place to place o r time

to time with d i f f e r e n t o r i g i n s a n d related

d e r i v a t i o n (gallinipper) S o m e t i m e s t h e s e

a r e literal translations f r o m m o d e r n lan­

g u a g e s (scorpions, escorpiones) o r usages (tarantulas) n o t c o m m o n to t h e r e g i o n

M i x t u r e s o f s y m p a t r i c l a n g u a g e s also o c c u r (sede [Spanish] + ocuilin [Náhuatl] = s e d e o -cuilin = silkworm) T h e only a t t e m p t to

s t a n d a r d i z e c o m m o n n a m e s h a s b e e n m a d e with pest species in English (Stoetzel 1989)

Most l a n g u a g e s have a b r o a d t e r m f o r insects a n d like animals, r o u g h l y equiva­

lent to t h e English, for e x a m p l e , " b u g "

("worm" o r " g r u b " ) : bicho (Spanish a n d

P o r t u g u e s e ) a n d ocuilin ( N á h u a t l )

References

AUDANT, A 1941 Identification des insectes d'Haiti par leur nom creóle Soc Hist Geogr

Haiti, Rev 12(42): 5 1 - 5 5 [Not seen.]

BAUCKE, O 1961 Os nomes comuns dos in­

sectos no Rio Grande do Sul Sec Agrie., Porto Alegre

BIEZANKO, C M., AND D LINK 1972 Nomes

populares dos Lepidópteros no Rio Grande

do Sul (Segundo Catalogo) Univ Fed Santa Maria, Bol Tec 4: 3 - 1 5

BRÜCHER, G 1942 Lista de algunos nombres vulgares de insectos Dept San Veg (Min

Agrie, Santiago) Bol 2(2): 120-125

DA SILVA, B R 1930-1934 Nomenclatura popular dos Lepidópteros do Distrito Federal and seus arredores Vols 1—5 O Campo, Rio

de Janeiro

DOUROJEANNI, M J 1965 Denominaciones ver­

naculares de insectos y algunos otros in­

vertebrados en la selva del Perú Rev Peruana Entomol 8: 131-137

GARCÍA, R J 1976 Nombre de algunos insectos

y otros invertebrados en "Quechua." Rev

Jíbaro-IHERING, R VON 1968 Dicionário dos animáis

do Brasil Ed Univ Brasilia, Sao Paulo

MONTE, O 1928 Os nomes vulgares dos in­

sectos de Brasil Almanak Agrie Brasil 1928:

228-289

MUNIZAGA, C , AND J HERRERA 1985 Notas

entomológicas de Socaire (Obtenidas durante

la Expedición Chileno-Alemana a Socaire, en mayo de 1957) Notas Centr Est Antropol., Univ Chile, 1: 3 - 1 3

ORDOÑO, C M 1982 Diccionario de zoología Náhuatl Ed Innovación, Mexico

PÉREZ D'ANGELLO, V 1966 Concordancia entre

los nombres vulgares y científicos de los insectos chilenos Mus Nac Hist Nat Not

Mens 10(119): 2 - 7

RIDE, W D L., C W SABROSKY, G BERN ARDÍ,

AND R V MELVILLE, eds 1985 International

code of zoological nomenclature 3d ed Intl

Trust Zool Nomen., London

STOETZEL, M B 1989 Common names of insects and related organisms Entmol Soc

Amer Lanham, Md

TASTEVIN, C 1923 Nomes de plantas e animaes

em Lingua Tupy Rev Mus Paulista 13: 6 8 7

-763

WELLING, E C 1958 Some Mayan names for certain Lepidoptera in the Yucatán penin­

sula J Lepidop Soc 12: 118

INSECTS AND HUMAN CULTURE

Aside from t h e i r i m p o r t a n c e as pests a n d

o u r a c a d e m i c i n t e r e s t in insects, these crea­

tures, s p i d e r s , a n d related a r t h r o p o d s have considerable influence in t h a t p o r t i o n of

tered t h r o u g h t h e r e m a i n d e r o f this b o o k

in t h e sections o n t h e v a r i o u s insects in­

volved; for Mexico, see M a c G r e g o r 1969.) Insects, s p i d e r s , c e n t i p e d e s , a n d scorpi­

ons a p p e a r in t h e Mayan Codices ( D r e s d e n ,

T r o - C o r t e s i a n u s , a n d Peresianus),

indicat-Figure 1.8 Decorative plates from modern

Peru prominently featuring ¡mages of the fly

(chuspi), revered in Incan times and a design

motif in Andean art today (Original, author's collection)

ing a n a p p r e c i a t i o n of their existence a n d their inclusion in c u l t u r a l events, such as rituals, c e r e m o n i e s , a n d d a n c e s T h e fa­

m o u s Nasca figures i n c l u d e a n i m m e n s e

s p i d e r (fig 1.9) Portions of s a m e a r e also stylized as glyphs h a v i n g linguistic signifi­cance (Tozzer a n d Allen 1910) I n t h e

e i g h t e e n t h century, it was believed t h a t a small, r e d insect (still u n i d e n t i f i e d b u t called "coya" in t h e O r i n o c o region) c a u s e d severe skin e r u p t i o n s ; its effects could only

be r e m e d i e d by c e r e m o n i o u s l y passing t h e body t h r o u g h a fire m a d e from a specific grass ("guayacán") ( K a m e n - K a y e 1979)

M a n y such c u r i o u s a c c o u n t s o f insects fill

t h e accounts o f early visitors a n d colonists

M o n t e z u m a ' s castle was built in w h a t is now p a r t of Mexico City, a n d U r u b a m b a ,

"plain of t h e insect" (uru = s p i d e r o r

evolved o n e s ; n o n e of t h e latter has yet

been o b s e r v e d in n a t u r e

Scientific n a m e s a r e p r o p e r l y p r o ­

n o u n c e d a c c o r d i n g to t h e rules of Latin,

but their way of b e i n g s p o k e n usually

varies a c c o r d i n g to t h e native accent of t h e

speaker T h i s s h o u l d b o t h e r n o o n e e x c e p t

Latin scholars, as l o n g as t h e n a m e is

u n d e r s t o o d

C o m m o n n a m e s , o r vulgates, a r e a p ­

plied to t h e insects a n d t h e i r relatives in

all Latin A m e r i c a n c o u n t r i e s Léxica have

been p u b l i s h e d for Chile ( B r ü c h e r 1942,

suffer from f r e q u e n t spelling a n d p r o n u n ­

ciation variations, p a r t i c u l a r l y in Brazil

a c c o r d i n g to n o consistent set of stan­

d a r d s , v a r y i n g f r o m place to place o r time

to time with d i f f e r e n t o r i g i n s a n d related

origin (scarabs o r escarabajos, from G r e e k

karabos) L a y m e n a n d c o u n t r y folk a r e likely

a r e literal translations from m o d e r n lan­

g u a g e s (scorpions, escorpiones) o r usages (tarantulas) n o t c o m m o n to t h e region

Mixtures of sympatric l a n g u a g e s also occur (sede [Spanish] + ocuilin [Náhuatl] = sedeo-cuilin = silkworm) T h e only a t t e m p t to

s t a n d a r d i z e c o m m o n n a m e s has b e e n m a d e with pest species in English (Stoetzel 1989)

Most l a n g u a g e s have a b r o a d t e r m for insects a n d like animals, r o u g h l y equiva­

lent to t h e English, for e x a m p l e , " b u g "

("worm" o r " g r u b " ) : bicho (Spanish a n d

P o r t u g u e s e ) a n d ocuilin ( N á h u a t l )

References

AUDANT, A 1941 Identification des insectes d'Haiti par leur nom creóle Soc Hist Geogr

Haiti, Rev 12(42): 5 1 - 5 5 [Not seen.]

BAUCKE, O 1961 Os nomes comuns dos in­

sectos no Rio Grande do Sul Sec Agrie, Porto Alegre

BIEZANKO, C M., AND D LINK 1972 Nomes

populares dos Lepidópteros no Rio Grande

do Sul (Segundo Catalogo) Univ Fed Santa Maria, Bol Tec 4: 3 - 1 5

BRÜCHER, G 1942 Lista de algunos nombres vulgares de insectos Dept San Veg (Min

Agrie, Santiago) Bol 2(2): 120-125

DA SILVA, B R 1930-1934 Nomenclatura popular dos Lepidópteros do Distrito Federal and seus arredores Vols 1-5 O Campo, Rio

de Janeiro

DOUROJEANNI, M J 1965 Denominaciones ver­

naculares de insectos y algunos otros in­

vertebrados en la selva del Perú Rev Peruana Entomol 8: J31-137

GARCÍA, R J 1976 Nombre de algunos insectos

y otros invertebrados en "Quechua." Rev

Jíbaro-IHERING, R VON 1968 Dicionário dos animáis

do Brasil Ed Univ Brasilia, Sao Paulo

42 GENERAL ENTOMOLOGY

MONTE O 1928 Os nomes vulgares dos in­

sectos de Brasil Almanak Agrie Brasil 1928:

028-289

MrxizAGA, C , AND J HERRERA 1985 Notas

entomológicas ¿ e Socaire (Obtenidas durante

la Expedición Chileno-Alemana a Socaire, en

mayo de 1957) Notas Centr Est Antropol.,

Univ Chile, 1: 3 - 1 3 ORDOÑO, C M 1982 Diccionario de zoología Náhuatl Ed Innovación, Mexico

PÉREZ D'ANGELLO, V 1966 Concordancia entre

los nombres vulgares y científicos de los insectos chilenos Mus Nac Hist Nat Not

Mens 10(119): 2 - 7

RIDE W D 1 , C W SABROSKY, G BERNARDI,

AND R V MELVILLE, eds 1985 International

code of zoological nomenclature 3d ed Int)

Trust Zool Nomen., London

STOETZEL, M B 1989 Common names of insects and related organisms Entmol Soc

Anier Lanham, Md

TASTEVIN, C 1923 Nomes de plantas e animaes

em Lingua Tupy Rev Mus Paulista 13: 6 8 7

-703

WELLING, E C 1958 Some Mayan names for certain I.epicloptera in the Yucatán penin­

sula J I.epidop Soc 12: 118

INSECTS AND HUMAN CULTURE

Aside from t h e i r i m p o r t a n c e as pests a n d our a c a d e m i c i n t e r e s t in insects, these crea­

tures, spiders, a n d related a r t h r o p o d s have considerable influence in t h a t p o r t i o n of

ples (Some o f t h e m o r e g e n e r a l a r e cited below; m a n y o t h e r specific cases a r e scat­

tered t h r o u g h t h e r e m a i n d e r of this book

in the sections o n t h e various insects in­

volved; for Mexico, see M a c G r e g o r 1969.) Insects, s p i d e r s , c e n t i p e d e s , a n d scorpi­

ons a p p e a r in t h e M a y a n Codices ( D r e s d e n , fro-Cortesianus, a n d Peresianus), indicat-

Figure 1.8 Decorative plates from modern

Peru prominently featuring images of the fly

(chuspi), revered in Incan times and a design

motif in Andean art today (Original, author's collection)

ing an a p p r e c i a t i o n of their existence a n d their inclusion in c u l t u r a l events, such as rituals, c e r e m o n i e s , a n d d a n c e s T h e fa­

m o u s Nasca figures include a n i m m e n s e spider (fig 1.9) Portions of s a m e a r e also stylized as glyphs h a v i n g linguistic signifi­cance (Tozzer a n d Allen 1910) In t h e

e i g h t e e n t h century, it was believed t h a t a small, red insect (still u n i d e n t i f i e d but called "coya" in t h e O r i n o c o region) caused severe skin e r u p t i o n s ; its effects could only

be r e m e d i e d by c e r e m o n i o u s l y p a s s i n g t h e body t h r o u g h a fire m a d e from a specific grass ("guayacán") ( K a m e n - K a y e 1979) Many such c u r i o u s a c c o u n t s of insects fill the accounts of early visitors a n d colonists

in t h e New World (Cowan 1865)

Insects have lent t h e i r n a m e s to m a n y places in Latin A m e r i c a A m o n g t h e better known a r e C h a p u l t e p e c , t h e "hill of t h e

g r a s s h o p p e r s " (chapulín = g r a s s h o p p e r + tepee = hill) w h e r e t h e Aztec E m p e r o r

M o n t e z u m a ' s castle was built in w h a t is now p a r t of Mexico City, a n d U r u b a m b a ,

"plain of t h e insect" (uru = s p i d e r o r

INSECTS AND HUMAN CULTURE 43

Figure 1.9 The spider was an eminent symbol in Peruvian cultures of prehistory It is displayed on a

grand scale among the Nasca figures in the southern desert

caterpillar + pampa = plain), the sacred

valley of the Incas near Cuzco in Peru

In modern times, insects symbolize nu­

merous ideas (fig 1.10), especially in litera­

ture and folklore (Lenko and Papavero 1979) Science fiction and fantasy novels often use the dangerous qualities of many types to instill horror or malevolence Su­

perstitions and fanciful stories attributing good or bad fortune to many insects, spiders, or the like, are believed by sectors

of the population, especially those in re­

mote or primitive areas (Hogue 1985)

T h e cultural use of insects is perhaps best developed among Indian tribes still surviving in many parts of Latin America (Berlin and Prance 1978; Hitchcock 1962;

Kevan 1983; Posey 1978, 1983) T h e study

of this aspect of cultural entomology is referred to as "ethnoentomology" (and includes some of the odd practical uses of

Figure 1.11 Image from the Codex Telleriano

Remensis of the Aztec deity, Itzpapálotl, in nature represented by wild silk moths of the genus

Rothschildia (Hand copy by Carlos spacher in Mariposas entre los Antiguos Mexi- canos, 1989; reproduced with author's permis­

Beutel-sion)

Figure 1.12 In a variation of the "toucandira

ritual" in which giant hunting ants of the genera

Dinoponera and Paraponera are used, a mat tied

with paper wasps is applied to the chest of this Roucouyenne Indian (French Guiana) to test his

courage (From H Davis, The Jungle and the Damned 1952, Duell, Sloan and Pearce, New

York; reproduced with permission)

insects, such as for food or medicine; see valuable insects, chap 3)

In many Amazonian Indian groups, insects were in the past and are still today venerated religiously, and they play central roles as deities or mythic figures (fig 1.11) The four guardians of the cardinal points

in Warao cosmology are social insects—two wasps, a bee, and a termite (Wilbert 1985) Ritual also incorporates insects, for exam­

ple, the giant hunting ants (Dinoponera) in

puberty ceremonies practiced by various Amazonian tribes (Liebrecht 1886; fig 1.12) Similarly, pain is endured from the stings of wasps whose nests are purposely molested as a part of rites of passage among the Gorotire-Kayapó in Brazil (Posey 1981)

Figure 1.13 Modern Peruvian Indian (Yagua)

necklace using beetle parts as main decorative

element (Original, author's collection)

Metallic beetle parts and even galls (Ber­

lin and Prance 1978) are used in body

ornamentation by Indians in all parts of

the region (fig 1.13) Insects, especially

musical species (crickets and katydids), lu­

minescent forms (headlight beetles), and

large beetles, and orthopterans are kept as

pets or curiosities Many species are eaten,

both for sustenance and as delicacies (fig

1.14)

References

BERLIN, B., AND G T PRANCE 1978 Insect galls

and human ornamentation: T h e

ethnobotan-ical significance of a new species of Licania

from Amazonas, Peru Biotropica 10: 81-86

COWAN, F 1865 Curious facts in the history of

insects Lippincott, Philadelphia

HITCHCOCK, S W 1962 Insects and Indians of

the Americas Entomol Soc Amer Bull 8(4):

Figure 1.14 A bottle of mezcal containing a

maguey worm (Comadla redtenbacheri,

Cossi-dae) as an extra treat for the drinker The beverage was important in ancient and modern Mexican culture The insect retains today its natural association with the plant and its product

(Los Angeles County Museum of Natural History collection)

KAMEN-KAYE, D 1979 A bug and a bonfire J

Ethnopharm 1: 103-110

KEVAN, Ó K M C E 1983 T h e place of grasshop­

pers and crickets in Amerindian cultures 2d Trien Meet Pan American Acrid Soc (Boze-man, Mont., 1979) Proc P 8-74c

LENKO, K., AND N PAPAVERO 1979 lnsetos no

folclore Conselho Estad Artes Cien Hu­

man., Sao Paulo

LIEBRECHT, F 1886 Tocandyrafestes Zeit

Ethnol 18: 350-352

MACGREGOR, R 1969 La representation des

insectes dans l'ancien Mexique giste25: 1-8

L'Entomolo-POSEY, D A 1978 Ethnoentomological survey

of Amerind groups in lowland Latin Amer­

ica Fia Entomol 61: 225-228

POSEY, D A 1981 Wasps, warriors and fearless

46 GENERAL ENTOMOLOGY

n c i r Fihnoentomology of the Kayapó Indi­

e s of central Brazil J Ethnobiol 1: 165-174

'SFY D A 1983 Ethnomethodology as an I'WiV guide to cultural systems: T h e case of the insects and the Kayapó Indians of Amazonia

Rev Brasil Zool 1: 135-144

)ZZER, A M-, AND G M ALLEN 1910 Animal

figures in the Maya Codices Harvard Univ, Peabody Mus., Pap., Amer Archaeol Ethnol 4: 273-372, pis 1-39

WILBERT J 1985 T h e house of the tailed kite: Warao myth and the art of think­

swallow-ing in images In G Urton, ed., Animal myths

and metaphors Univ Utah, Salt Lake City

INSECTS AND HUMAN CULTURE 47

HUFFAKER, C B., AND R L RABB, eds 1984

Ecological entomology Wiley, New York

PRICE, P W 1984 Insect ecology 2d ed Wiley,

New York

WALTER, H 1979 Vegetation of the earth and

ecological systems of the geo-sphere 2d ed

Springer, New York Translated from 3d rev

a n d relief, o r its p h y s i o g r a p h y , which p r o ­

vides f o o t h o l d s for t h e very existence a n d

b r o a d areas within Latin A m e r i c a which

d e t e r m i n e in a most e l e m e n t a l way t h e

d i s t r i b u t i o n of insects P h y s i o g r a p h i c s u b d i ­visions in Latin A m e r i c a have b e e n o u t ­lined a c c o r d i n g to various s c h e m e s A simplified version is p r e s e n t e d below (Fig

2.1); it is modified f r o m S a u e r (1950) a n d Sick (1969)

References

SAUER, C O 1950 Geography of South Amer­

ica, Handbk So Amer Indians 6: 319-344

SICK, W D 1969 Geographic substance In E J

Fittkau, J lilies, H Klinge, G H Schwabe, and H Sioli, eds., Biogeography and ecology

in South America 2: 449-474 Junk, T h e Hague

Climate and Medium

Each climatic factor e x e r t s its critical ef­

fects in a variety of ways: t e m p e r a t u r e as freezing point, highs, lows, m e a n s , r a n g e s , heat, cold, daily fluctuations; m o i s t u r e as rainfall, dew, fog, clouds (and, of c o u r s e ,

by d e t e r m i n i n g t h e f u n d a m e n t a l

life-s u p p o r t i n g m e d i a , aquatic v e r life-s u life-s t e r r e life-s ­trial); sunlight as day a n d n i g h t , s h a d e , illumination, r a d i a t i o n , a n d p h o t o p e r i o d All act o v e r l o n g time p e r i o d s as w e a t h e r

Figure 2.1 MAJOR PHYSIOGRAPHIC AREAS OF LATIN AMERICA (from Sauer 1950 and Sick

1969) MIDDLE AMERICA: 1 Mexican Highlands; 2 Isthmian America (lowland Mexico, Central America); 3 West Indies (Greater and Lesser Antillean Islands); 4 Bahamas; SOUTH AMER­ICA: 5 Pacific Coastal plain; 6 Andes (including Caribbean Borderlands and Bolivian Altiplano);

7 Amazon Basin; 8 Orinoco Basin; 9 Guiana Highlands; 10 Brazilian Highlands (including Brazilian Coastal Mountains) and Mato Grosso, Plateau of Paraná; 11 Llanos de Mamoré; 12 Paraná-Paraguay Depression; 13 Gran Chaco; 14 Pampas; 15 Patagonia; INSULAR AMERICA:

16 Oceanic islands (Pacific: Galapagos and Revillagigedo Archipelagos, Cocos Island, Easter Island,

vegetation flourishes (especially o n t h e

lomas; see Special H a b i t a t s , below) T h e

effects of El N i ñ o o n coastal insect p o p u l a ­

across t h e A m a z o n Basin in o d d years,

usually d u r i n g early J u l y (Días Frios d e San

R e s p o n s e s of insects to these stresses

may b e d r a m a t i c o r subtle b u t a r e as yet

virtually u n s t u d i e d in Latin A m e r i c a T h e

impact of such s u d d e n w e a t h e r c h a n g e s

on t h e s e small, e c t o t h e r m i c c r e a t u r e s surely m u s t b e intense, especially o n

t e m p e r a t u r e - s e n s i t i v e species I n t e r f e r ­ence with r e p r o d u c t i o n a n d p o p u l a t i o n

"die-offs" s h o u l d be e x p e c t e d

T h e effects of climate a r e felt most

i m m e d i a t e l y in t h e a t m o s p h e r e T h e y a r e manifest equally b u t m o r e slowly i n t h e aquatic m e d i u m , in s t a n d i n g (lentic) waters ( p o n d s , lakes, etc.), in m o v i n g (lotic) waters (streams, rivers) (Fittkau 1 9 6 4 ; M a c a n

1962, 1974), a n d in t h e soil ( K ü h n e l t 1961)

References

BEINGOLEA, O D 1987a El fenómeno "El Niño" 1982-83 y algunos insectos-plaga en la costa peruana Rev Peruana Entomol 28:

55-57

BEINGOLEA, O D 19876 La langosta Schislocerca interrita en la costa norte del Perú, durante

1983 Rev Peruana Entomol 28: 3 5 - 4 0

KÓPPEN, W., AND R GEIGER 1931-1934

Hand-buch der Klimalologie Vols 1 —4 traeger, Berlin

Born-KÜHNELT, W 1961 Soil biology with special

reference to the animal kingdom Faber 8c

Faber, London

FITTKAU, E J 1964 Remarks on limnology of central-Amazon rain-forest streams Int

Verh Limnol., Verh 15: 1092-1096

MACAN, T T 1962 T h e ecology of aquatic insects Ann Rev Entomol 7: 261-288

MACAN, T T 1974 Freshwater ecology 2d ed

Wiley, New York

PHILANDER, S G H 1990 El Niño, La Niña and the Southern Oscillation Academic, San Diego

RATISBONA, L R 1976 T h e climate of Brazil, hi

W Schwerdtfeger, ed., Climates of Central and South America, world survey of climatol­

ogy 12: 219-293 Elsevier, Amsterdam

Vegetation Zones

Physicochemical e n v i r o n m e n t a l factors n o t only act directly o n insect life f o r m s b u t influence t h e m indirectly t h r o u g h t h e

k i n d s of p l a n t g r o w t h they allow (biotic factors) G r o u p i n g s of plants a d a p t e d to a

p a r t i c u l a r set of soil a n d climatic c o n d i t i o n s delimit b r o a d e n v i r o n m e n t s f o r insects

50 ECOLOGY

Many types of v e g e t a t i o n a r e p r e s e n t in Latin A m e r i c a , a n d t h e s e a r e classified according to v a r i o u s systems (e.g., B e a r d

1944, G r a h a m 1 9 7 3 , H u e c k a n d Seibert

1 9 7 2' S a u e r 1950, W e b e r 1969) T h e s e systems a r e only regionally applicable b e ­cause they often define u n i t s in t e r m s of specific plant taxa p r e s e n t A universal scheme, globally a p p l i c a b l e , is H o l d r i d g e ' s (1967, 1982), which c o m b i n e s t h e effects of elevation, l a t i t u d e , rainfall, a n d t e m p e r a ­ture to define v e g e t a t i o n a l f o r m a t i o n s , in­

d e p e n d e n t of floristic e l e m e n t s

It is very a p p a r e n t t h a t t h e n a t u r e of vegetation plays a p r i m a r y r o l e in d e t e r m i n ­ing t h e N e o t r o p i c a l insect f a u n a in e a c h physiographic a r e a T h e special r i c h n e s s of the A m a z o n Basin is a g o o d case in point

C o n t r a r y to f o r m e r ideas, t h e region's vege­

tation h a s n o t existed c o n t i n u o u s l y u n ­

c h a n g e d for t e n s of millions of years b u t h a s varied c o n s i d e r a b l y f r o m n e a r d e s e r t t o lush forest in r e c e n t geologic p e r i o d s , p a r ­ticularly d u r i n g t h e Pleistocene A g e , d u r ­ing a l t e r n a t i n g arid a n d h u m i d c o n d i t i o n s

In t h e d r i e r p h a s e s , m o i s t u r e - r e q u i r i n g vegetation s h r u n k greatly a n d f r a g m e n t e d into forest p a t c h e s w h e r e rainfall persisted which was a d e q u a t e for t h e i r survival ("ref­

uge theory," H a f f e r 1 9 8 2 ; b u t see E n d l e r 1982)

T h i s disjunction i n t o forest islands iso­

lated from e a c h o t h e r by g r a s s l a n d o r even desertlike p l a n t cover d i v i d e d m a n y for­

merly c o n t i n u o u s p o p u l a t i o n s a n d led to their evolution i n t o n e w species Wet phases, such as t h e w o r l d is n o w e x p e r i e n c ­ing, allowed t h e p a t c h e s to e x p a n d again and t h e gaps b e t w e e n t h e m to close B u t

evidences of t h e f o r m e r islands, o r refugia,

are still p r e s e n t as c o n c e n t r a t i o n s of e n ­demics A m o n g insects, this is s h o w n espe­

cially well by butterflies ( B r o w n 1982) a n d

a m o n g a r a c h n i d s by s c o r p i o n s ( L o u r e n c o 1986)

T h i s h e t e r o g e n e i t y in A m a z o n i a n forest and wet forests in o t h e r a r e a s partly e x ­plains t h e latest a n d o n e of t h e most

BROWN, JR., K S 1982 Paleoecology and re­

gional patterns of evolution in Neotropical

forest butterflies In G T Prance, ed., Biologi­

cal diversification in the tropics Columbia Univ Press, New York Pp 255-308

ENDLER, J A 1982 Pleistocene forest refuges:

Fact or fancy In G T Prance, ed., Biological

diversification in the tropics Columbia Univ Press, New York Pp 641-657

GRAHAM, A., ed 1973 Vegetation and vegeta­tional history of northern Latin America Elsevier, Amsterdam

HAFFER, J 1982 General aspects of the Refuge

Theory In G L Prance, ed., Biological diver­

sification in the tropics Columbia Univ Press, New York Pp 6 - 2 4

HOLDRIDGE, L R 1967 Life Zone ecology Trop Sci Ctr., San José, Costa Rica

HOLDRIDGE, L R 1982 Ecología basada en zonas de vida Insto, lnteramer Coop Agrie San José, Costa Rica

HUECK, K., AND P SEIBERT 1972

Vegeta-tionskarte von Südamerika (Mapa de la vegetación de America del sur) Fischer, Stuttgart

LOURENC.O, W R 1986 Diversité de la faune scorpionique de la region amazonienne; cen­tres d'endémisme; nouvel appui á la théorie des refuges forestiers du Pleistocene Amazo-

n i a n a 9 : 559-580

SAUER, C O 1950 Geography of South Amer­ica Handbk So Amer Indians 6: 319-344

SIMPSON, B B., AND J HAFFER 1978 Speciation

patterns in the Amazonian forest biota Ann Rev Ecol Syst 9 : 4 9 7 - 5 1 8

WEBER, H 1969 Zur natürlichen

Vegeta-tionsgliederung von Südamerika In E J

Fittkau, J lilies, H Klinge, G H Schwabe, and H Sioli, eds., Biogeography and ecology

in South America 2:475-518 Junk, T h e Hague

Artificial Environments

T h e f o r e g o i n g discussion h a s b e e n con­

c e r n e d with n a t u r a l o r original c o n d i t i o n s

a n d p a t t e r n s of native flora a n d f a u n a Since c o m i n g to t h e s o u t h e r n l a n d s o f t h e

GEOGRAPHY 51

New W o r l d 2 0 , 0 0 0 to 5 0 , 0 0 0 years a g o ,

h u m a n s h a v e modified t h e original life

zones to v a r y i n g d e g r e e s (Kiinkel 1963),

even c r e a t i n g l a r g e tracts of essentially

new, "artificial life z o n e s " Such a r e t h e

cities, f a r m s , a n d vast g r a s s l a n d s for cattle

cultivated species t h a t only slowly r e t u r n e d

to climax status t h r o u g h successional

KÜNKEL, G 1963 Vegetationszerstorung und

Bodenerosion in Lateinamerika Arch

w h e r e t h e d e t e r m i n a n t s of life specifically affect a n o r g a n i s m is its m i c r o h a b i t a t Ex­

a m p l e s a r e infinite in n u m b e r ; t h e i n t e r i o r

of a d e c a y i n g log o r water-filled c u p of a

b r o m e l i a d a r e e x a m p l e s T h e vertical distri­

b u t i o n of different species of m o s q u i t o e s (Bates 1944) a n d butterflies ( P a p a g e o r g i s 1975) in forests is e v i d e n c e of t h e effects of subtle e n v i r o n m e n t a l d e t e r m i n a n t s T h e existence of so m a n y m i c r o h a b i t a t s partly accounts for t h e vast diversity of t h e in­

g r a p h i c features (caves, lakes), a r e m a c r o habitats Still g r e a t e r g r o u p i n g s a n d l a r g e r

-e x p a n s -e s of -e a r t h f o r m i n g grossly r-ecogniz­

able m a c r o h a b i t a t s a r e called life zones o r

b i o m e s (e.g., tropical forest, d e s e r t s ) T h e statuses of Latin A m e r i c a n insects in sev­

eral special habitats a r e discussed below

Special Habitats

T h e g r e a t n u m b e r of n a t u r a l N e o t r o p i c a l insect m a c r o - a n d m i c r o h a b i t a t s m a k e s it impossible to discuss m o r e t h a n a few of t h e most distinctive, peculiar, a n d i m p o r t a n t

T h e y f o r m p a r t i c u l a r places for insects whose s t r u c t u r e a n d activities m a y b e very

d i f f e r e n t f r o m those of t h e o t h e r insects in

t h e s u r r o u n d i n g g e n e r a l e n v i r o n m e n t Most insects s t u d i e d in artificial habitats

a r e injurious species a n d a r e discussed in

C h a p t e r 3, Practical E n t o m o l o g y A few investigations have focused o n o t h e r fau-nal e l e m e n t s of p l a n t a t i o n s (Young 1986a, 19866), habitations, a n d like a r e a s u n d e r

h u m a n m a n a g e m e n t

References

BATES, M 1944 Observations on the distribu­

tion of diurnal mosquitoes in a tropical forest

of u p p e r - s t o r y life T h e c a n o p y could b e reached by skilled native climbers, b u t they carried n o scientific e x p e r t i s e aloft, a n d their activities h a d to b e d i r e c t e d ineffec­

tively by t h e i r e a r t h b o u n d e m p l o y e r s

O t h e r a p p r o a c h e s , still r e m o t e o n e s , have been to elevate v a r i o u s kinds of t r a p s to catch s o m e of t h e f a u n a o r k n o c k it d o w n with quick-acting, b i o d e g r a d a b l e insecti­

cides (Erwin 1983)

G r e a t e r i m p r o v e m e n t in access c a m e with t h e c o n s t r u c t i o n of a r b o r e a l l a d d e r s and p l a t f o r m s ( P o r t e r a n d DeFoliart 1981), towers (a f a m o u s o n e o n B a r r o C o l o r a d o Island, P a n a m a ) , o r elevated causeways, from which o b s e r v a t i o n s a n d collections could b e m a d e H o w e v e r , these offer very limited mobility a n d a r e costly to c o n s t r u c t and m a i n t a i n

Lately, s o m e practical m e a n s h a v e b e e n found n o t only to m o v e a b o u t a n d obtain specimens f r o m this c o m p l e x r e a l m b u t even to carry o u t e x t e n d e d studies within it (Mitchell 1982) Even scientists themselves are now able to g o i n t o t h e canopy, u s i n g

m o u n t a i n e e r i n g t e c h n i q u e s (Perry 1980,

1984; Perry a n d Williams 1981) U n f o r t u ­nately, few e n t o m o l o g i s t s a r e t r a i n e d in

b o t h t h e a c a d e m i c a n d athletic aspects of this d e m a n d i n g , a l t h o u g h highly r e w a r d ­ing, a p p r o a c h

T h e results of these efforts, a l t h o u g h

References

ERWIN, T L 1983 Beetles and other insects of tropical forest canopies at Manaus, Brazil, sampled by insecticidal fogging techniques

In S L Sutton, T C Whitmore, and A C

Chadwick, eds., Tropical rain forest: Ecology and management Blackwell, Oxford Pp 59-75

HINGSTON, R W 1932 A naturalist in the

Guiana forest Longmans Green, New York MITCHELL, A W 1982 Reaching the rain forest roof (A handbook on techniques of access and study in the canopy) Leeds Phil Lit Soc Leeds, Eng

PERRY, D R 1980 1 probe the jungle's last frontier Int Wildlife 10: 5 - 1 1

PERRY, D R 1984 T h e canopy of the tropical rain forest Sci Amer 251: 138-147

PERRY, D R., AND J WILLIAMS 1981 T h e

tropical rain forest canopy: A method provid­ing total access Biotropica 13: 283-285

PORTER, C LL, AND G R DEFOLIART 1981 T h e

man-biting activity of phlebotomine sand flies (Díptera: Psychodidae) in a tropical wet forest environment in Colombia Arq Zool Sao Paulo 30: 81-158

Amazon Inundation Forests

Vast e x p a n s e s b o r d e r i n g t h e A m a z o n a n d its major t r i b u t a r i e s a r e a n n u a l l y flooded

to a d e p t h of several m e t e r s for p e r i o d s of five to six m o n t h s o r m o r e T h e s e forests

(called igapó locally) h a r b o r an insect f a u n a

especially a d a p t e d to t h e stresses o f alter­nately rising a n d r e c e d i n g waters

T h e terrestrial species t h a t live in these

forests a r e n u m e r o u s a n d very diverse

spiracles situated o n s h o r t stalks to aid

b r e a t h i n g while they a r e in t h e water O n l y

ADIS, J 1977 Programa mínimo para analises

de ecosistemas: Artrópodos terrestres em

florestas inundáveis da Amazonia central

Acta Amazónica 7: 223-229

A D I S , J., AND V MAHNERT 1985 On the natural

history and ecology of Pseudoscorpions

(Arachnida) from an Amazonian blackwater

inundation forest Amazoniana 9: 297-314

A D I S , J., V MAHNERT, J W DE MORÁIS, AND

J M GOMES 1988 Adaptation of an Amazo­

nian pseudoscorpion (Arachnida) from dry­

land forests to inundation forests Ecology

6 9 : 2 8 7 - 2 9 1

ERWIN, T L., AND J A D I S 1982 Amazonian

inundation forests: Their role as short-term

refuges and generators of species richness

and taxon pulses In G T Prance, ed., Biologi­

cal diversification in the tropics Columbia

Univ Press, New York Pp 3 5 8 - 3 7 1

Soil and Ground Litter

T h e soil a n d its overlying layer of o r g a n i c litter constitute t h e habitat of a g r e a t

m a n y small t o very small a r t h r o p o d s (Kiihnelt 1961) D o m i n a n t a m o n g these

a r e ants, springtails, s p i d e r s , psocids,

t h r i p s , cryptostigmatic mites, a n d beetles (Penny et al 1978), b u t l a r g e r insects a r e s o m e t i m e s also f o u n d h e r e (Young 1983) It is i n c r e d i b l e t h a t such

micro-a r t h r o p o d s m micro-a y c o m p r i s e micro-almost 5 0 p e r ­cent of t h e total soil a n d litter biota in t h e

c e n t r a l A m a z o n forest, a n t s a n d t e r m i t e s

a l o n e f o r m i n g a b o u t 6 0 p e r c e n t of this (Fittkau a n d Klinge 1973) Most o c c u p y

t h e u p p e r 5 to 8 c e n t i m e t e r s of t h e soil

T h e c o m p o s i t i o n o f this f a u n a a n d its ecology have b e e n t h e subject of m a n y studies in t h e N e o t r o p i c s , especially in for­

ests (e.g., Williams 1941, Deboutteville a n d

R a p o p o r t 1 9 6 2 - 1 9 6 8 , L i e b e r m a n a n d Dock 1982) w h e r e m o i s t u r e seems to be t h e major factor d e t e r m i n i n g t h e seasonal dis­

DANTAS, M., AND H O R SCHUBART 1980

Correlacáo dos indices de agregacáo de Acari

e Collembola com 4 fatores ambientáis numa pastagem de terra firme da Amazonia Acta Amazónica 10: 771-774

DEBOUTTEVILLE, C D., AND E RAPOPORT, eds

1962-1968 Biologie de 1'Amérique Australe

Etudes sur la faune de sol; 2: Etudes sur la faune du sol; 3: Etudes sur la faune du sol +

54 ECOLOGY

Documents biogéographiques; 4, Documents biog et ecologiques Ed Cen Nat Rech Sci., Paris

FITTKAU, E J., AND H KLINGE 1973 On

biomass and trophic structure of the central Amazonian rain forest ecosystem Biotropica

5 : 2 - 1 4 KÜHNELT, W 1961 Soil biology with special reference to the animal kingdom Faber &

Faber, London

LEVINGS, S C , AND D M WINDSOR 1984 Litter

moisture content as a determinant of litter arthropod distribution and abundance dur­

ing the dry season on Barro Colorado Island, Panama Biotropica 16: 125-131

LIEBERMAN, S., AND C F DOCK 1982 Analysis

of the leaf litter arthropod fauna of a lowland tropical evergreen forest site (La Selva, Costa Rica) Rev Biol Trop 30: 2 7 - 3 4

PEARSON, D L., AND J A DERR 1986 Seasonal

patterns of lowland forest floor arthropod abundance in southeastern Perú Biotropica 18: 244-256

PENNY, N D., J R ARIAS, AND H O R

SCHUBART 1978 Tendencias populacionais

de fauna de Coleópteros d o solo sob floresta

de terra firme na Amazonia Acta Amazónica 8: 259-265

STRICKLAND, A H 1945 A survey of the

arthropod soil and litter fauna of some forest reserves and cacao estates in Trinidad, British West Indies J Anim Ecol 14: 1-11

WILLIAMS, E C 1941 An ecological study of the floor fauna of the Panamanian rain forest Chicago Acad Sci Bull 6: 63-124

WILLIS, E D 1976 Seasonal changes in the invertebrate litter fauna on Barro Colorado Island, Panamá Rev Brasil Biol 36: 6 4 3 -

657

YOUNG, A M 1983 Patterns of distribution and abundance in small samples of litter-inhabiting Orthoptera in some Costa Rican cacao plantations New York Entomol Soc J

9 1 : 3 1 2 - 3 2 7

Black Water Lakes and Rivers

Some lowland tropical river basins c o n t a i n tributaries a n d l a n d l o c k e d basins (oxbow

lakes, cochas) with tea-colored w a t e r t h a t in

the d e p t h s a p p e a r s black T h e s e a r e distin­

guished from so-called white waters n o t only by their color b u t by physical a n d chemical p r o p e r t i e s W h i t e waters (actually

a milky chocolate color) a r e n u t r i e n t rich, neutral to slightly alkaline, a n d t u r b i d

Black waters usually flow from n u t r i e n t poor, sandy soils a n d t h u s a r e low in minerals b u t a r e acidic a n d m a y c o n t a i n high c o n c e n t r a t i o n s o f o r g a n i c c o m p o u n d s (tannic acids, phenolics) leached f r o m vege­tation a n d toxic to insects T h e w a t e r a n d

a n d water mites (Tundisi et al 1979), c a n

be very n u m e r o u s , even in t h e most heavily

c h a r g e d water

T h e G u i a n a Shield of n o r t h e r n S o u t h

A m e r i c a is a l a r g e black w a t e r a r e a a n d is

n o t o r i o u s for t h e p o o r n e s s of its p r o d u c t i v ­ity O t h e r similar such r e g i o n s a r e f o u n d in

t h e Brazilian H i g h l a n d s a n d o n t h e Yuca­tán Peninsula

Clear waters ( g r e e n i s h t o clear) a r e also

r e c o g n i z e d b u t a r e biologically similar t o black water

References

FITTKAU, E J 1971 Distribution and ecology of Amazonian chironomids (Diptera) Can Ento­mol 103: 407-413

JANZEN, D H 1974 Tropical blackwater rivers, animals and mast fruiting by the Diptero-carpaceae Biotropica 6: 69—103

TUNDISI, J G., A M P MARTINS, AND T

MATSU-MURA 1979 Estudos ecológicos preliminares

em sistemas aquáticos em Aripuaná Acta Amazónica 9: 311-315

Caves

Insects a n d m a n y r e l a t e d t e r r e s t r i a l a r t h r o ­

p o d s of diverse g r o u p s a r e t r u e troglobites (obligate cavernicoles, i.e., animals n a r ­rowly a n d specifically a d a p t e d for life d e e p

in caves; C u l v e r 1982, H o f f m a n n et al 1986) However, tropical caves a r e usually

d o m i n a t e d by species classed as t r o g l o philes, which also live in n o n c a v e habitats

-S o m e a r e o m n i v o r e s , b u t they a r e m o r e

n o r m a l l y specialized f o r p a r t i c u l a r foods

a n d a r e of two basic types: s c a v e n g e r s a n d

ECOSYSTEMS 55

uals A m o n g these a r e t h e terrestrial

i s o p o d s (Trichorhina), millipedes

(Eurhino-cricus), c o c k r o a c h e s {Periplaneta, Blaberus),

tineid m o t h s , d u n g beetles (Ataenius), a n d

C a v e insects often exhibit m o r p h o l o g i ­

cal a d a p t a t i o n s to life in t h e d a r k (Dessen

et al 1980), i n c l u d i n g eyes r e d u c e d o r

a b s e n t , lack of i n t e g u m e n t a r y p i g m e n t a ­

tion, r e d u c t i o n a n d loss o f wings, a n d

greatly e l o n g a t e d , highly sensitive a p p e n d ­

1980) Recent work, yet u n p u b l i s h e d , has

f o u n d over thirty species of eyeless cave

a n d soil a r t h r o p o d s in volcanic caves in t h e

DESSEN, E M B., V R ESTON, M S SILVA,

M T TEMPERINI-BECK, AND E TRAJANO

1980 Levantamento preliminar de fauna de cavernas de algumas regioes do Brasil Cien

Cult 3 2 : 4 1 4 - 7 2 5 GNASPINI, N R 1989 Análise comparativa da fauna associada a depósitos de guano de morcegos cavernícolas no Brasil: Primeira Approximafáo Rev Brasil Entomol 32:

ORGHIDAN, T , A NÚÑEZ JIMÉNEZ, V DECON,

S NEGREA, AND N V BAYES 1973-1983

Résultats des expeditions biospéleologique cubano-Roumaines á Cuba Vols 1-4 Ed

Academiei, Bucharest, Republicii Socialiste Romania

PECK, S B 1974 T h e invertebrate fauna of tropical American caves Pt II: Puerto Rico,

an ecological and zoogeographic approach

Biotropica 6: 14—31

PECK, S B 1975 T h e invertebrate fauna of tropical American caves Pt I l l : Jamaica, an introduction Int.J Speleol 7: 303-326

PECK, S B 1977 Recent studies on the inverte­

brate fauna and ecology of sub-tropical and tropical American caves 6th Intl Cong

Speleol Proc 5: 185-193

PECK, S B 1981a Zoogeography of inverte­

brate cave faunas in southwestern Puerto Rico Nati Speleol Soc Bull 43: 70-79

PECK, S B 19816 Community composition and zoogeography of the invertebrate cave fauna

of Barbados Fla Entomol 64: 519-527

REDDELL, J R 1971 A review of the cavernicole fauna of Mexico, Guatemala, and Belize

Texas Mem Mus Austin Bull 27: 1-327

STRINATI, P 1971 Recherches logiques en Amerique du Sud Ann Speleol

biospéleo-26: 439-450

Lomas

T h e low coastal d e s e r t hills of c e n t r a l P e r u provide o n e of t h e m o s t u n u s u a l habitats for insects in t h e N e o t r o p i c ( D o g g e r a n d Risco 1970) It is s p e c u l a t e d t h a t , histori­

cally, t h e l o m a s w e r e p a r t of a l a r g e r

c h a p a r r a l b i o m e t h a t o n c e e x t e n d e d a l o n g the e n t i r e w e s t e r n slopes of t h e A n d e s (Péfaur 1978) I n this z o n e of e x t r e m e general aridity, wetness in t h e f o r m of r a i n comes only at m u l t i y e a r intervals, d e c a d e s

or m o r e At t h e s e times, explosions of plant life o c c u r o n t h e o t h e r w i s e p a r c h e d hills, a n d they b e c o m e g r e e n islands in t h e bleak d e s e r t Normally, only t h e r e g u l a r

a r t h r o p o d s , s o m e species of which a r e known from n o w h e r e else (Aguilar 1964)

T h e m o r e i m p o r t a n t g r o u p s a r e s p i d e r s (Aguilar, P a c h e c o , a n d Silva 1987), mites, springtails, wax insects, beetles (especially

T e n e b r i o n i d a e ) , flies, a n d a n t s (Aguilar 1981), which a r e m o s t a b u n d a n t a n d n u ­merous in k i n d s in t h e d a m p e r u p p e r elevations S o m e special f o r m s a r e wing­

less sticklike f o r m s ("palitos vivientes d e Lima"), i n c l u d i n g two walkingsticks, a

j u m p i n g stick (Proscopiidae), a n d assassin

bugs ( R e d u v i i d a e ) O n e walkingstick thra minúscula) is o m n i v o r o u s b u t dies with

(Libe-the failing p l a n t s at t h e e n d of t h e d a m p

season T h e o t h e r (Bostra scabrinota) feeds

on o n e p l a n t b u t can c h a n g e its color to match seasonal c h a n g e s a n d is p r e s e n t

y e a r - r o u n d (Aguilar 1970) A n u n e x ­pected e l e m e n t is t h e w a t e r m e a s u r e r

(Bacillometra woytkowskii, H y d r o m e t r i d a e ) ,

a h e t e r o p t e r o u s insect n o r m a l l y associated with bodies of fresh w a t e r (Aguilar, Oyeyama, a n d A g u i l a r 1987) Its p r e s e n c e

is associated with t h e h i g h h u m i d i t y of t h e lomas in t h e winter

References

AGUILAR, P G 1964 Especies de artrópodos registrados en las lomas de los alrededores de Lima Rev Peruana Entomol 7: 9 3 - 9 5 AGUILAR, P G 1970 Los "palitos vivientes de Lima." 1: Phasmatidae de las lomas Rev Peruana Entomol 13: 1—8

AGUILAR, P G 1981 Fauna desértico-costera Peruana V i l : Apreciaciones sobre diversidad

de invertebrados en la costa central Rev Peruana Entomol 24: 127-132

AGUILAR, P G., F OYEYAMA, AND Z P AGUILAR

1987 Los "palitos vivientes de Lima." 111: Un Hydrometridae de las lomas costeras Rev Peruana Entomol 28: 8 9 - 9 2

AGUILAR, P G., V R PACHECO, AND R SILVA

1987 Fauna desértico-costera peruana VIII: Arañas de las lomas Zapalla!, Lima (nota preliminar) Rev Peruana Entomol 29: 99—

103

DOGGER, J., AND S H Risco 1970 La fauna insectil de las lomas de Trujillo, Estudio del cerro "Campana." Bol Tec Circ Entomol Norte (Lambayeque) 1(2): 1-5

PÉFAUR, J E 1978 Composition and structure

of communities in the lomas of southern Peru Ph.D diss., Univ Kansas, Lawrence

a n d p o n d s i n l a n d a n d at t h e ocean's m a r ­gin T h e r e a s o n s a p p a r e n t l y lie in t h e fact

t h a t they a r r i v e d o n t h e scene l o n g after their o t h e r i n v e r t e b r a t e p r e d e c e s s o r s h a d locked u p all t h e ecological niches T h e r e

s t r i d e r s , c h a p 7) Closer to s h o r e , t h e

n u m b e r a n d kinds of m a r i n e insects greatly increases T h e r e o n e finds t h e

m a r i n e m i d g e s ( C h i r o n o m i d a e ) T h e s e live

o n rocky s h o r e s a n d h a v e larvae t h a t a r e

Several kinds of lice a r e parasitic o n

s e a g o i n g m a m m a l s , seals a n d sea lions

-saline lake types, t h a t h a v e shifted to t h e

similar habitats at t h e sea m a r g i n s , e s p e ­

cially salt m a r s h e s a n d m a n g r o v e s A major­

f e e d i n g e i t h e r directly o n this m a t e r i a l (kelp

fly larvae, Fucellia; see c h a p 1 1 ;

m a n life impossible in seaside a r e a s Several

w a t e r b o a t m e n species (especially

Tricho-corixa reticulata) live in highly saline s h o r e

pools (Davis 1966) a n d h a v e even b e e n

DAVIS, C C 1966 Notes on the ecology and

reproduction of Trichocorixa reticulata in a

Jamaican salt-water pool Ecology 47: 850—

852

EVANS, W G 1968 Some intertidal insects from

western Mexico Pan-Pacific Entomol 44:

b o u n d i n g o v e r b o u l d e r s a n d c r a s h i n g i n t o

f o a m - c o v e r e d pools, t o r r e n t i a l s t r e a m s d e ­scend t h r o u g h g o r g e s a n d n a r r o w can­

yons, over h a r d , rocky b e d s , b e f o r e r e a c h ­ing t h e lower, gentle slopes

Fast, cold w a t e r offers a refugial n i c h e to

m a n y insects, c o m p a r a t i v e l y free f r o m ver­

t e b r a t e p r e d a t i o n b e c a u s e few such l a r g e animals can live in such a n i n h o s p i t a b l e

m e d i u m E x c e p t i o n s a r e a few h a r d y insecti­

v o r o u s types, such as t o r r e n t d u c k s , d i p ­

p e r s , a n d a few fish ( i n t r o d u c e d t r o u t a n d possibly s o m e a s t r o b l e p h i d catfishes) As a result, s o m e very a n c i e n t r e p r e s e n t a t i v e s of several aquatic g r o u p s h a v e p e r s i s t e d for geologic ages as t o r r e n t i c o l o u s ( r h e o p h i -lous) relicts T h e s e i n c l u d e e n t i r e families, most p r o m i n e n t l y t h e n e t - w i n g e d m i d g e s ( B l e p h a r i c e r i d a e ) a n d various beetles (Elmi-dae, D r y o p i d a e , P s e p h e n i d a e ) O t h e r simi­

larly a d a p t e d taxa a r e t h e larval stages of

lance-winged m o t h flies (Maruina,

Psy-c h o d i d a e ) a n d blaPsy-ckflies (Simuliidae) a n d

m a n y species of T r i c h o p t e r a (especially in

t h e family H y d r o b i o s i d a e ) , H e m i p t e r a phocricos, N a u c o r i d a e ) , P l e c o p t e r a (Arau- canioperla, G r i p o p t e r y g i d a e ) , w a t e r m i d g e s ( C h i r o n o m i d a e ) , a n d E p h e m e r o p t e r a (Epe- orus, H e p t a g e n i i d a e )

[Cry-E x t r e m e s t r u c t u r a l a n d physiological ad­

a p t a t i o n s have evolved in t h e s e f o r m s in

r e s p o n s e to t h e s t r o n g selective p r e s s u r e s of fast c u r r e n t a n d s m o o t h s u b s t r a t e s T h e s e include holdfast s t r u c t u r e s (ventral suck­

ers, claws), s t r e a m l i n i n g , a n d p l a s t r o n respi­

r a t i o n T h e latter m a k e s u s e of t h e function

of air films a n d pockets as "physical gills." A

r e q u i r e m e n t for p r o p e r function of this

58 ECOLOGY

system is cold, clean, o x y g e n - r i c h water

T h e p r e s e n c e of these insects t h e r e f o r e indicates h e a l t h y s t r e a m c o n d i t i o n s

Food habits for t h e relatively passive grazers a n d p r e d a t o r s d o n o t r e q u i r e r a p i d

m o v e m e n t A d u l t e m e r g e n c e is often "ex­

plosive." To avoid d r o w n i n g , t h e i m a g o rises to t h e s u r f a c e in a b u b b l e a n d takes wing i m m e d i a t e l y after c o n t a c t i n g t h e air

T o r r e n t i c o l o u s insects a r e n o t well known in Latin A m e r i c a S o m e f a m o u s early studies w e r e m a d e o n D i p t e r a in

s o u t h e a s t e r n Brazil by Fritz Müller (1879, 1895) a n d A d o l f o L u t z (1930) N e w species are c o m m o n l y d i s c o v e r e d in t h e habitat, especially in r e m o t e m o u n t a i n s

References

LUTZ, A 1930 Biología das aguas torrenciais e encachoeiradas Soc Biol Montevideo, Arch

suppl 1: 114-120

MÜLLER, F 1879 A metamorphose de uni

insecto díptero Mus Nac Rio de Janeiro, Arch 5/6: 4 7 - 8 5 , pis 4 - 7

MÜLLER, F 1895 Contributions towards the history of a new form of larvae of Psychodidae (Diptera) from Brazil Entomol Soc London, Trans 1895: 479-482, pis 1 0 - 1 1

Tank Plants

Some N e o t r o p i c a l plants h a v e p a r t s of their a n a t o m y d e v e l o p e d i n t o c u p - s h a p e d water-holding s t r u c t u r e s ( p h y t o t e l m a t a )

(Frank a n d L o u n i b o s 1983) a n d a r e r e ­

ferred to as r e s e r v o i r o r t a n k p l a n t s T h e y are of several types T h e best k n o w n a r e the b r o m e l i a d s (Bromeliaceae) (Gómez 1977) w h o s e w a t e r a c c u m u l a t i o n s p r o v i d e

a h o m e for small a q u a t i c a n i m a l s T h i s microcosm was first s t u d i e d c o m p r e h e n ­sively as a n ecosystem by Picado (1913),

who d i s t i n g u i s h e d b e t w e e n t h e a q u a t i c mi­

lieu ( " a q u a r i u m " ) a n d t e r r e s t r i a l p o r t i o n

("terrarium")

T h e a q u a r i u m consists of a spacious central c u p a n d e x p a n d e d lateral leaf bases

that collect r a i n w a t e r L a r g e p l a n t s m a y

store a liter o r m o r e a n d m a y have a c u p 5 to

10 c e n t i m e t e r s , in d i a m e t e r a n d equally

d e e p T h e s e reservoirs of w a t e r p r o v i d e habitats suitable for t h e d e v e l o p m e n t of

(Chironornus), p u n k i e s (Bezzia), damselflies

( P s e u d o s t i g m a t i d a e ) , beetles ( H e l o d i d a e ) ,

a n d water mites T h e detailed ecology of this fraction h a s b e e n s t u d i e d by m a n y entomologists, i n c l u d i n g Laessle (1961) (See t h e b i b l i o g r a p h y of b r o m e l i a d a n d

p i t c h e r plant r e s e r v o i r p l a n t e n t o m o l o g y by Fish a n d B e a v e r [1978].)

D e t r i t u s collects also in t h e lateral bracts,

a n d a special kind of a r b o r e a l soil is c r e ­ated which is like a " t e r r a r i u m " to a n o t h e r

g r o u p of insects H e r e a r e f o u n d s p i d e r s , carabid beetles, ants, isopods, millipedes, mites, springtails, a n d o t h e r t e r r e s t r i a l forms (Murillo et al 1983) A few insects actually feed o n t h e leaves of b r o m e l i a d s

a n d f o r m yet a n o t h e r guild in association with these plants ( B e u t e l s p a c h e r 1972)

A n o t h e r category of N e o t r o p i c a l t a n k plants a r e t h e insectivorous " p i t c h e r p l a n t s "

of t h e family S a r r a c e n i a c e a e I n t h e s e ,

d e e p , u r n - s h a p e d leaves h a v e evolved to hold fluids t h a t n o r m a l l y kill a n d digest hapless insects t h a t fall into t h e m However,

i m m a t u r e s of s o m e insect species, mostly mosquitoes of t h e tribe Sabethini, a r e im­

m u n e to t h e corrosive action of these c h e m i ­cals a n d actually d e v e l o p n o r m a l l y in this very peculiar aquatic m i c r o h a b i t I n t h e

N e o t r o p i c s , p i t c h e r plants of only t h e g e n u s

Heliamphora a r e k n o w n , f o u n d in t h e

V e n e z u e l a n - G u i a n a n r e g i o n T h e y a r e

k n o w n to be o c c u p i e d by Wyeomyia m o s q u i ­ toes of t h e s u b g e n u s Zinzala (Zavortink

1985), b u t their o t h e r o c c u p a n t s a r e n o t

s t u d i e d Flower bracts, especially those of t h e

ECOSYSTEMS 59

beetles in older, less v o l u m i n o u s bracts with

small a m o u n t s o f water T h e most c o m m o n

fly larvae a r e of p o m a c e flies

(Droso-philidae), h o v e r flies ( S y r p h i d a e ,

Cope-stylum), a n d soldier flies (Stratiomyiidae,

Merosargus) T h e beetles a r e p r i m a r i l y

s c a v e n g i n g w a t e r beetles ( H y d r o p h i l i d a e ,

Gillisius), leaf beetles ( C h r y s o m e l i d a e ,

His-p i n a e , CeHis-phaloleia), a n d r o v e beetles

of water At least five g e n e r a a r e r e p r e ­

s e n t e d : Wyeomyia, Triehoprosopon,

Toxorhyn-chites, Culex, a n d Sabethes

BEUTELSPACHER, C R 1972 Some observations

on the Lepidoptera of bromeliads J Lepidop

Soc 26: 133-137

DOWNS, W G., AND C S PITTENDRIGH 1946

Bromeliad malaria in Trinidad, British West

Indies Amer J Trop Med Hyg 26: 4 6 - 6 6

FISH, D., AND R A BEAVER 1978 A bibliogra­

phy of the aquatic fauna inhabiting bromeli­

ads (Bromeliaceae) and pitcher plants thaceae and Sarraceniaceae) Florida Anti-mosq Assoc, Proc 49: 11-19

(Nepen-FRANK, J H., AND L P LOUNIBOS, eds 1983

Phytotelmata, terrestrial plants as hosts for aquatic insect communities Plexus, Medford,

N.J

GÓMEZ, L D 1977 La biota bromelícola ex­

cepto anfibios y reptiles Hist Nat Costa Rica 1:45-62

LAESSLE, A M 1961 A micro-limnological

study of Jamaican bromeliads Ecology 42:

499-517

MURILLO, R M.,J G PALACIOS, J M LABOUGLE,

E M HENTSCHEL, J E LLÓRENTE, K LUNA,

P ROJAS, AND S ZAMUDIO 1983 Variación

estacional de la entomofauna asociada a

Tillandsia spp en una zona de transición

biótica Southwest Entomol 8: 292-302

PICADO, C 1913 Les broméliacées epiphytes considérée comme milieu biologique Bull

Scient France Belgique 47: 215-360

SEIFERT, R P 1975 Clumps of Heliconia

inflorescences as ecological islands Ecology 56: 1416-1422

SEIFERT, R P 1980 Mosquito fauna of Heliconia áurea J Anim Ecol 49: 687-697

SEIFERT, R P 1981 Principal components analy­

sis of biogeographic patterns among Heliconia

insect communities New York Entomol Soc,

J 89: 109-122

SEIFERT, R P 1982 Neotropical Heliconia insect

communities Quart Rev Biol 57: 1-28

SEIFERT, R P., AND F H SEIFERT 1976a Natu­

ral History of insects living in inflorescences

of two species of Heliconia New York

Entomol Soc.,J 84: 233-242

SEIFERT, R P., AND F H SEIFERT 19766 A

community matrix analysis oí Heliconia insect

communities Amer Nat 110: 4 6 1 - 4 8 3

SEIFERT, R P., AND F H SEIFERT 1979 A

Heliconia insect community in a Venezuelan

cloud forest Ecology 60: 462-467

ZAVORTINK, T J 1985 Zinzala, a new subgenus

of Wyeomyia with two new species from

pitcher-plants in Venezuela (Diptera, dae, Sabethini) Wasmann J Biol 43: 46-59

Culici-Miscellaneous Special Habitats

A few a d d i t i o n a l i m p o r t a n t special habitats

t h a t have b e e n i g n o r e d o r received mini­

mal a t t e n t i o n with r e g a r d to t h e i r e n t o m o faunal aspects in Latin A m e r i c a a r e t h e following

-1 Hot, mineral springs D e Oliveira (1954)

n o t e d e p h y d r i d flies in t h e h o t effluent

of a geyser at El T a t i o in Chile S o m e

g e n e r a l collections f r o m T e r m a s d e Chillan in t h e s a m e c o u n t r y h a v e b e e n identified by Ruiz a n d S t u a r d o (1935)

2 Inland salt lakes and playas, such as t h e

i m m e n s e salares of t h e Bolivian alti­

plano (Salar d e Coipasa, L a g o d e Poopó, Salar d e U y u n i ) A few insects of

h y p e r s a l i n e w a t e r s h a v e b e e n d e ­scribed, notably m o s q u i t o e s (Bach-

h a r d - b o t t o m s t r e a m s (lilies 1964; C a m ­pos et al 1984; Patrick a n d collabs

1966; T u r c o t t e a n d H a r p e r 1982a, 19826) A d a p t i v e strategies w e r e stud­

ied o n s t r e a m i n v e r t e b r a t e s , mostly in­

sects in t h e A r g e n t i n e R i o N e g r o , by

W a i s ( 1 9 8 5 )

4 Alpine ponds and lakes Available a r e

only t h e r e p o r t s o f Robeck e t al (1980) for m i s c e l l a n e o u s A n d e a n habitats a n d Gilson (1939) f o r L a k e Titicaca

5 Bogs N o p u b l i s h e d o b s e r v a t i o n s

6 Deserts, b o t h local a n d l a r g e , as t h e

great S o n o r a n o r A t a c a m a N o a p p a r ­ent c o m p r e h e n s i v e o b s e r v a t i o n s

7 Sand dunes, b o t h coastal a n d i n l a n d

f o u n d o n n e w volcanoes, b o t h at sea

a n d o n t h e c o n t i n e n t s I n b o t h cases,

t h e principles of island b i o g e o g r a p h y

a r e applicable

10 Guianan tepuis T h e flat-topped j u n g l e

mesas of t h e G u i a n a Shield a r e ac­

k n o w l e d g e d islands s u p p o r t i n g m a n y

e n d e m i c biotic e l e m e n t s T h e y h a v e

b e e n s t u d i e d mostly by botanists, b u t

s o m e u n i q u e insects h a v e b e e n col­lected a t o p M o u n t R o r a i m a a n d C e r r o Neblina ( S p a n g l e r 1 9 8 1 , 1985; W a t e r -

h o u s e 1900)

11 Lakes and rivers Limnological studies,

particularly of bodies of fresh w a t e r

l a r g e r t h a n s t r e a m s , a r e n u m e r o u s , b u t

in these, m o r e a t t e n t i o n is usually paid

to t h e m a c r o f a u n a t h a n to insects

S o m e works have c o n c e n t r a t e d o n t h e latter (Wais 1984)

12 Land crab burrows T h e b u r r o w s o f

tropical coastal land c r a b s s u p p o r t a

c o m p l e x ecosystem, i n c l u d i n g m a n y insects a n d related a r t h r o p o d s ( B r i g h t

a n d H o g u e 1972), most o f which h a v e aquatic life stages in t h e w a t e r t h a t

a c c u m u l a t e s within t h e m f r o m rainfall

a n d g r o u n d w a t e r s e e p a g e

13 Lava tubes N o a p p a r e n t o b s e r v a t i o n s

Also r e m a i n i n g to b e e l u c i d a t e d a r e t h e entomological characteristics of t h e g r e a t

n u m b e r of vegetation types t h r o u g h o u t

t h e r e g i o n , for e x a m p l e , c e r r a d o , p a l m forests, caatinga, c l o u d forest, m a n g r o v e s , grassland, p u n a , p á r a m o ( B e r n a l 1985), alpine, a n d s a v a n n a

T h e f o r e g o i n g a r e all fruitful fields o f investigation for f u t u r e s t u d e n t s of ecologi­cal a n d b i o g e o g r a p h i c entomology

References

BACHMANN, A O 1979 Notas para una monografía de las Corixidae Argentinas (In-

secta, Heteroptera) Acta Zool Lilloana 35:

305-350

BACHMANN, A O., AND O H CASAL 1963

Mosquitos Argentinos que crian en aguas

salobres y saladas Soc Entomol Argentina,

Rev 25: 2 1 - 2 7

BERNAL, A 1985 Estudio comparativo de la

artropofauna de un bosque alto-andino y de

un pajonal paramuno en la región de

Monserrate (Cund.) In H Sturm and

O Rangel, Ecología de los paramos andinos:

Una visión preliminar integrada Univ Nac

Colombia (lnst Cien Nat., Mus Nat Hist.,

Bibl José Jerónimo Triana no 9) Bogotá Pp

225-260

BRIGHT, D B., AND C L HOGUE 1972 A

synopsis of the burrowing land crabs of the

world and list of their arthropod symbionts

and burrow associates Los Angeles Co Mus

Nat Hist., Contrib Sci 220: 1-58

CAMPOS, H., J ARENAS, C JARA, T GONSER, AND

R PRINS 1984 Macrozoobentos y fauna

lótica d e las aguas limnéticas de Chiloé y

Aysén continentales (Chile) Medio Ambiente

7: 5 2 - 6 4

DE OLIVEIRA, S J 1954 Contribuicáo para o

conhecimento do género "Dimecoeniá"

Cres-son, 1916 1: "Dimecoenia lenti" sp n encon­

trado no Chile (Díptera, Ephydridae) Rev

Brasil Biol 14: 187-194

DE OLIVEIRA, S J 1958 Contribuicáo para o

conhecimento d o género "Dimenia" Cresson,

1916 IV: Descricáo da larva e do pupário de

"Dimecoenia grumanni" Oliveira, 1954

(Diptera, Ephydridae) Rev Brasil Biol 18: 1 6 7

-169

GILSON, H C 1939 T h e Percy Sladen Trust

Expedition to Lake Titicaca in 1937 I: De­

scription of the expedition Linnean Soc

London, Trans 1: 1—20

ILLIES, J 1964 T h e invertebrate fauna of the

Huallaga, a Peruvian River, from the sources

down to Tingo María Intl Verein Theor

Angew Limnol., Verh 15: 1077-1083

PALACIOS-VARGAS, J 1985 Microartrópodos del

Popocatepetl (aspectos ecológicos y

biogeo-gráficos de los ácaros oribátidos e insectos

colémbolos) Ph.D diss., Univ Nac Aut Méx­

ico, Mexico City

PATRICK, R., AND COLLABORATORS 1966 T h e

Catherwood Foundation Peruvian-Amazon

Expedition: Limnological and systematic stud­

ies Acad Nat Sci Philadelphia, Monogr 14:

1-495

ROBECK, S S., L BERNER, O S FLINT, J R ,

N NIESER, AND P J SPANGLER 1980 Results

of the Catherwood Bolivian-Peruvian alti­

plano expedition Pt 1, Aquatic insects except Diptera Acad Nat Sci Philadelphia Proc 132: 176-217

Ruiz, E, AND C STUARDO O 1935 Insectos colectados en las Termas de Chillan Rev

Chilena Hist Nat 39: 313-322

SPANGLER, P J 1981 New and interesting water beetles from Mt Roraima and Ptari-tepui, Venezuela (Coleóptera: Dytiscidae and Hy-drophilidae) Aquat Ins 3: 1-11

SPANGLER, P J 1985 A new genus and species

of riffle beetle, Neblinagena prima, from the

Venezuelan tepui, Cerro de la Neblina (Coleóptera, Elmidae, Larinae) Entomol

Soc Wash., Proc 87: 538-544

TURCOTTE, P., AND P P HARPER 1982a T h e

macroinvertebrate fauna of a small Andean stream Freshwater Biol 12: 411-419

TURCOTTE, P., AND P P HARPER 19826 Drift

patterns in a high andean stream biologia89: 141-151

Hydro-WAIS, 1 R 1984 Two Patagonian basins—

Negro (Argentina) and Valdivia (Chile)—as habitats for Plecoptera Ann Limnol 20:

115-122

WAIS, I R 1985 Strategies adaptatives aux eaux courantes des invertébrés d u bassin du fleuve Negro, Patagonia, Argentina Int

Verein Limnol., Verh 22: 2167-2172

WATERHOUSE, C O 1900 Report on a collec­

tion made by Messrs E V McConnell and J J

Quelch at Mt Roraima in British Guyana

Linnean Soc London, Trans 8: 74—76

HISTORICAL BIOGEOGRAPHY

T h e historical p e r s p e c t i v e in zoogeog­

r a p h y p r o v i d e s insights i n t o t h e o r i g i n s o f species a n d f a u n a s a n d their c h a n g e s

t h r o u g h time relative to geologic events (Fittkau et al 1969) Insects m a k e excellent

m a t e r i a l for such studies b e c a u s e of t h e very long history of m a n y taxa a n d t h e variety of types available t o test h y p o t h e s e s

of all kinds (Gressitt 1974, M u n r o e 1965)

A g r o u p m a y b e f o u n d which will fulfill almost a n y given set of c o n d i t i o n s f o r any geologic p e r i o d For e x a m p l e , u p l a n d , sed­

e n t a r y types with limited vagility (such as

"dispersalist" a n d "vicariance" schools of biogeographic t h o u g h t (Ferris 1980) to explain t h e speciation p r o c e s s in evolving organisms Actually, b o t h m e c h a n i s m s may cause t h e b r a n c h i n g of phyletic lines and a r e p a r t o f a m o d e r n , unified t h e o r y

of b i o g e o g r a p h y (Pielou 1979, B r o w n a n d Gibson 1983)

Because insects h a v e b e e n o n e a r t h for a very long t i m e , at least since t h e m i d d l e of the Paleozoic e r a , c o n t i n e n t a l c o n n e c t i o n s and disjunctions (tectonics) (Dietz a n d Holden 1970, M a r v i n 1 9 7 3 , S m i t h et al

1981) a r e major vicariant events t h a t h a v e affected their evolution a n d dispersal (Car-bonell 1977: 1 5 5 - 1 6 1 ) I n Latin A m e r i c a , the tectonic d e v e l o p m e n t of t h e C a r i b b e a n and isthmian r e g i o n s s e e m s to be m u c h

m o r e c o m p l i c a t e d ( B o n i n i e t al 1984, D u r ­ham 1985, R o s e n 1985) t h a n t h a t o f t h e South A m e r i c a n p o r t i o n s ( J e n k s 1956,

H a r r i n g t o n 1962) with c o n s e q u e n t p r o b ­lems in e x p l a i n i n g t h e o r i g i n s of o r g a n i s m s there ( L i e b h e r r 1988, W o o d s 1989)

T h e o r i g i n of m a n y g r o u p s o n G o n d wanaland, t h e g r e a t s o u t h e r n c o n t i n e n t that was c o m p o s e d o f w h a t is n o w S o u t h America, Africa, A n t a r c t i c a , Australia, a n d India, is still e v i d e n t in t h e restricted occur­

-rences of t h e i r d e s c e n d a n t s in those a r e a s and in t h e s o u t h e r n m o s t p o r t i o n s o f S o u t h America t o d a y (e.g., w a t e r m i d g e s [ B r u n -din 1966, 1967]; s e e Keast 1973 for o t h e r insect e x a m p l e s of these so-called amphinotic o r a u s t r a l disjunctive distribu­

tions) Close affinities of s o m e e a s t e r n Brazilian insects with West African species,

such as a m o n g Schistocerca g r a s s h o p p e r s

(Carbonell 1977:169), t h e a m b l y p y g i d g e ­

nus Phrynus ( = T a r a n t u l a ) ( Q u i n t e r o 1983), t h e psocid g e n e r a Belaphapsocus a n d Notiopsocus ( N e w 1987), a n d t h e t e r m i t e

A m a z o n River system was p r o b a b l y c o n ­

t i n u o u s d u r i n g t h e early Mesozoic with t h e

N i g e r River, a n d t h e m a i n flow was west­

w a r d to t h e Pacific O c e a n B u t with t h e relatively r e c e n t (Miocene) uplift o f t h e

A n d e s , t h e flow was d a m m e d a n d a n

e n o r m o u s lake f o r m e d at their foot As t h e plates s e p a r a t e d , t h e w e s t e r n p o r t i o n o f

t h e river r e v e r s e d its flow a n d c a m e t o

e m p t y into t h e Atlantic A n extensive coastal plain was also c r e a t e d a l o n g t h e west side of t h e C o r d i l l e r a f r o m ejection o f volcanic m a t e r i a l a n d pluvial o u t w a s h

o n e r e a s o n for t h e basin's i n c r e d i b l e spe­cies richness

O n a m u c h m o r e limited scale, local

n a t u r a l disasters, such as volcanic e r u p t i o n s

a n d h u r r i c a n e s , take a toll o n insect life a n d may actually cause t h e extinction o f small

p o p u l a t i o n s o r even very r e g i o n a l species

a n d constitute vicariant events, a l t h o u g h such c o n s e q u e n c e s h a v e yet to b e d o c u ­

m e n t e d , especially those of low-density for­est species (Elton 1975) C h a n g e s in t h e

c o u r s e o f rivers, a n especially c o m m o n

o c c u r r e n c e in m e a n d e r i n g lowland d r a i n ­ages, such as t h e A m a z o n , c a n b r e a k u p

c o n t i n u o u s p o p u l a t i o n s a n d halt g e n e flow sufficiently t o c r e a t e new entities

Dispersalist e x a m p l e s o f t h e effects o f geology o n t h e history of insect life c o m e

f r o m t h e oceanic islands scattered o n t h e

HISTORICAL BIOGEOGRAPHY 63