I

atas do simpósio

SÔBRE A

biota amazônica

Vol. 5: Zoologia

Na Capa: Pavâozinho-do-Pará, Eurypygas h. he-

llas (Palias), ave endêmica da região

amazônica. (Foto de Moacyr Leão).

ATAS DO SIMPÓSIO

SÔBRE A

BIOTA AMAZÔNICA

Vol. 5: zoologia

Belém, Pará, Brasil, Junho 6-11, 1966

EDITOR: HERMAN LENT

Publicado pelo

CONSELHO NACIONAL DE PESQUISAS

RIO DE JANEIRO, GB

196 7

Illljl

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15

APRESENTAÇAO

De 6 a 11 de junho de 1966, na cidade de Belém, Estado do Pará,

Brasil, foi realizado o Simpósio sôbre a Biota Amazônica, organizado

pela Associação de Biologia Tropical, com a colaboração do Conselho

Nacional de Pesquisas do Brasil, tendo José Cândido de Melo Carvalho

como Presidente Executivo.

O Simpósio homenageava especialmente o Museu Paraense “Emilio

Goeldi” que comemorava seu 100.° aniversário.

Ao se iniciarem os trabalhos, achavam-se inscritos no Simpósio 16

países representados por 97 instituições, 256 pesquisadores inscritos para

apresentação de trabalhos que perfaziam um total de 22 conferências

e 198 contribuições originais. Associaram-se como observadores, até êsse

dia, 103 pessoas. Nos dias que se seguiram, até o encerramento, o total

geral de frequência dos inscritos foi a 611 pessoas. As contribuições ori-

ginais também aumentaram para 227.

Resolvemos editar estas Atas em 7 volumes, cada qual correspon-

dendo a uma das seções do Simpósio : Geociências, Antropologia, Limno-

logia, Botânica, Zoologia, Patologia e Conservação da Natureza e Recur-

sos Naturais; serão todos publicados pelo Conselho Nacional de Pesquisas

do Brasil, que assumiu a responsabilidade global da edição, da mesma

forma como promoveu a realização e apoiou a execução do Simpósio.

Em relação ao Programa do Simpósio distribuído na ocasião e,

ainda, ao próprio desenrolar das reuniões de cada Seção, as Atas não

incluem necessàriamente todos os trabalhos, retirados que foram alguns

por motivos vários.

Êste quinto volume corresponde à Seção V (Zoologia) que teve

como Coordenadores: Paulo E. Vanzolini (Departamento de Zoologia,

São Paulo) e Jesus S. Moure (Universidade do Paraná); consta de um

total de 604 páginas e 257 figuras no texto, e divulga 31 trabalhos, dos

quais três conferências. O índice do volume aparece a seguir pela ordem

alfabética do sobrenome dos autores, primeiro as conferências e depois

as comunicações.

Herman Lent

Setembro, 1967

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ÍNDICE DO VOLUME 5: ZOOLOGIA

Págs-

Dodson, Calaway H.

Relationships between pollinators and orchid flowers (Con-

ferência) 1

Stanley Rand, A.

Predator-prey interactions and the evolution of aspect diversity

(Conferência) 73

Vanzolini, P. E.

Problems and programs in Amazonian zoology (Conferência) ... 85

Beck, Ludwig

Die Bodenfauna des neotropischen Regenwaldes 97

Bokermann, Werner C. A.

Notas sôbre a distribuição de Bufo granulosus Spix, 1824 na

Amazônia e descrição de uma subespécie nova (Amphibia,

Bufonidae) 103

Bücherl, Wolfgang

Escorpiões, aranhas e escolopendromorfos da Amazônia 111

Cerqueira, N. L.

Simuliidae da Amazônia. III: Sôbre o gênero Simulium Latreille,

1802 (Diptera, Nematocera) 127

Cunha, Osvaldo Rodrigues da

Lacertílios da Amazônia. III: O gênero Arthrosaura Boulenger,

1885 (Lacertilia, Teiidae) 141

Dressler, Robert L.

Why do Euglossine bees visit orchid flowers? 171

Gonçalves, Cincinnato Rory

As formigas cortadeiras da Amazônia, dos gêneros Atta Fabr. e

Acromyrmex Mayr (Hym., Formicidae) 181

Gorenz, August M.

Períodos de secreção nectarífera na região de Belém, Pará . . . 203

Gorenz, August M.

Resumo de 5 anos de estudos sôbre apicultura no Baixo Amazonas 207

Handley, Jr., Charles O.

Bats of the canopy of an Amazonian forest 211

Hoge, A. R.

Serpentes do Território Federal do Amapá 217

1 ágs-

Iide, Paulo

Estudo sôbre uma nova espécie amazônica do gênero Cyphomyia

Wiedemann (Diptera, Stratiomyidae) 225

Jimbo, Sachiko & Schwassmann, Horst O.

Feeding behavior and the daily emergence pattern of Artibeus

jamaicensis Leach (Chiroptera, Phyllostomidae) 239

Kerr, Warwick E.; Sakagami, Shôichi F.; Zucchi, Ronaldo; Portugal-

Araujo, Virgilio & Camargo, João Maria F. de

Observações sôbre a arquitetura dos ninhos e comportamento de

algumas espécies de abelhas sem ferrão das vizinhanças de

Manaus, Amazonas (Hymenoptera, Apoidea) 255

Lemos de Castro, Alceu

Isópodos terrestres da Amazônia brasileira (Isopoda, Oniscoidea) 311

Lent, Herman & Jurberg, José

Revisão dos Piratinae americanos. III: As espécies do gênero

Tydiães Stal, com um estudo sôbre a genitália (Hemiptera,

Reduviidae) 337

Machado-Allison, C. E.

Sobre algunos ectoparásitos de mamíferos de la Biota Amazônica 365

Moure, J. S.

Descrição de algumas espécies de Euglossinae (Hym., Apoidea) 373

Moure, J. S.

A check-list of the known Euglossine bees (Hymenoptera, Apidae) 395

Oliveira, S. J. de

Nôvo gênero de Chironomidae da Amazônia (Insecta, Diptera) 417

Orejas-Miranda, Braulio R.

El género Leptotyphlops en la región amazônica 421

Peracchi, Adriano Lúcio

Lista remissiva dos Cléridas da Amazônia (Coleoptera, Cleridae) 443

Piza Jr., S. de Toledo

Uma nova espécie de Pseudomiopteriginae do Brasil 473

Righi, Gilberto

Descrição de Rhinodrilus priollii sp. n., Glossoscolecidae da

Amazônia, com bibliografia dos Oligochaeta terricola da região 475

Schiapelli, Rita D- & Pikelin, Berta S. Gerschman de

Estúdio sistemático comparativo de los generos Theraphosa

Walck., 1805; Lasiodora C. L. Koch, 1851 y Sericopelma Ausserer,

1875 (Araneae, Theraphosidae) 481

Sick, Helmut

Rios e enchentes na Amazônia como obstáculo para a avifauna 495

Thomé, José Willibaldo

A ocorrência de Veronicellidae na Amazônia brasileira, com des-

crição de duas espécies novas (Soleolifera, Gastropoda) 521

Vulcano, M. A. & Pereira, F. S.

Sinopse dos Passalidae e Scarabaeidae s. str. da Região Amazônica

(Insecta, Coleoptera) 533

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Atas do Simpósio sôbre a Biota Amazônica

Vol. 5 (Zoologia): 1-72 — 1967

RELATIONSHIPS BETWEEN POLLINATORS AND

ORCHID FLOWERS

CALAWAY H. DODSON

University of Miami, Coral Gables, Florida, USA.

(With 42 text-figures)

Throughout the literature the

remarkable characteristics, modifi-

cations and colorations of orchid

flowers have been discussed at

length. Seldom, however, has there

been serious speculation supported

by field observations, concerning

the reasons behind those often exo-

tic shapes and colors in such ka-

leidoscopic abundance. Why do

members of Cochlioda, Sphronitis

and certain species of Laelia and

Denãrcbium have extraordinarily

brilliantly colored flowers? Why do

the genera Catasetum, Cycnoches,

Stanhopea, Coryanthes, Gongora,

and others, have flowers which

almost defy description in their

strange complexities? Why do some

orchid flowers look like insects?

Why do we encounter elaborate

traps, springing or balanced lips,

tassels and ribbon-like sepals and

petals in the orchids? And finally,

why do orchid flowers produce so

many different odors; some spicey,

some sweet and some* incredibly

foul? Some authors have felt that

the rich variety in the orchid fami-

ly can be attributed to “the whims

of nature”. Others suggest “physi-

ology” or “genetics”. Darwin wrote

a book on the subject in 1862 and

correctly suggested that these cha-

racterists are adaptations to the

pollinators of the flowers.

Darwin unfortunately only had

limited information concerning

pollination of tropical orchids

though he did gather considerable

data on the temperate orchids of

Europe. The really striking floral

mechanisms are found, for the

most part, in tropical orchids, and

Darwin’s book contains presump-

tions and intuitions as to the me-

chanisms of many of these orchids

derived from careful study of the

flowers themselves. Nevertheless,

he had no way of knowing the truly

1 — 37 145

Atas do Simpósio sobre a Biota Amazônica

incredible relationships between

these weird flowers and their pol-

linators.

After Darwin there was a lapse

of nearly 100 years before intense

interest in the orchid flowers and

their pollinators revived. The reviv-

al was due primarily to similar

studies in other plant families. To

be sure, excellent work in temper-

ate orchids in Europe and Aus-

trália led to the discovery of pseu-

docopulation in Ophrys (Correvan

& Pouyanne, 1916) and Cryptosty-

lis (Coleman, 1928-1932). In Eu-

rope a nurríber of botanists contin-

ued to speculate about the mean-

ings of the complexities of the

tropical orchid flowers but without

observational data. Porsch (1955)

observed some pollination of Tro-

pical American orchids but missed

the significance of much of what

he saw. Ridley (1894) described

pollination of some Asian orchids

but only became interested in this

phase toward the end of his long

and productive career.

In recent years new and concert-

ed interest in pollination dynam-

ics has developed. Allen (1951,

1952, 1954) observed pollination of

several Central American orchids.

Kullenberg (1956, 1961) studied

pseudo-copulation in Oprhys. Vo-

gel (1959, 1963) published a study

of the pollination of temperate

orchids of South África. He also

worked on tropical orchids and

their pollinators in the Western

Hemisphere, and has published a

fine resume of pollinator behavior

in regard to Catasetum, Stanhopea

and allied groups. My work (Dod-

son & Frymire, 1961 a & b and

Dodson, 1962 a & b, 1965 a & b,

1966) has contributed field obser-

vations in quantity and detail in

an effort to make speculation

about the role of pollinators in the

evolution of the orchid species less

haphazard.

The new effort has demonstrat-

ed a criticai need for knowledge of

behavior and life histories of the

pollinators, for only with under-

standing of why they visit the flow-

ers can we complete the picture. A

simple list of which vector visits

which species of orchid tells us

much about the classification and

evolution of species in the family.

However, we find a large number

of orchids which do not attract

their pollinators on a simple “food

platter” basis, but tend to deceive

them in many ways. In order to

understand evolution and specia-

tion in those orchids an intensive

study of the behavior and the life

cycles of their pollinators was

needed and is now under way by

several workers.

THE POLLINATORS

Though bees appear to be the

basic and most abundant pollina-

tors of orchids, a number of other

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Volume 5 (Zoologia)

3

insects and a few specialized birds

are also functional.

BEES AND WASPS

Bees tend to be strong fliers, and

to have good Vision and perception

of odors. They can force their way

into stout flowers and discriminate

well between different kinds of

flowers. Logically they could act

as the primary agents in the pol-

lination of the orchid flower that

was already strongly “pollinator

oriented” in its early development.

Wasps do not fly as well as bees

and tend to be less effective as pol-

linators of specialized flowers.

Bee and wasp flowers — Flow-

ers which are adapted for pollina-

tion by bees and wasps, in the broad

sense, are called melittophilous.

Both bees and wasps belong to the

large and advanced order Hyme-

noptera and we usually refer to the

whole group of flowers which they

visit as “bee-flowers”. Some of the

primary characteristics of “bee-

-flowers” are that they are open

during the day when most bees and

wasps fly, they tend to produce

agreeable odors and bright colors

to attract the insects, they tend

to be zygomorphic with the devel-

opment of strong landing plat-

forms, they tend to have néctar

guides in the form of colored lines,

running into the depths of the

flower and they tend to have con-

cealed nectaries.

A curious characteristic of bees

is that of “constancy”, in which a

single bee or a whole colony of bees

will concentrate on a single species

of flowering plant in collecting

néctar and pollen. It is a learned

reaction which changes as the flow-

er supply changes but not all bees

demonstrate it. A quite different

reaction associates certain bees

with certain species of flowering

plants instinctively. This reaction

runs the gammut from polytropic

(visiting many kinds of flowers)

to oligotropic (visiting a limited

number of kinds) to monotropic

(where a 1-1 relationship exists

between a single species of bee and

a single species of flowering plant) .

These phenomena have consider-

able importance for flowering

plants adapted to certain species of

bees in a given area will be discuss-

ed below.

Bee and wasp-pollinated orchids

have the basic characteristics of

the “bee-flower” in general. They

usually have one of the petals well

developed into a large, extended

blade upon which the pollinator

lands. The basal portions of the lip

are usually formed into a tunnel

with the column forming the upper

side. The bee enters the tunnel to

get at the nectary and in backing

out some of the stigmatic liquid

may be rubbed on its dorsal sur-

face. As the bee backs further the

pollinia become attached to the

Atas do Simpósio sôbre a Biota Amazônica

sticky material and are carried to

the next flower. In more advanced

orchids the pollinia are fitted with

caudicles or may be attached to a

sticky pad, the viscidium, which

becomes detached from part of the

stigma and sticks to the pollinator.

Bee-pollinated orchids also have

colors in the range of the bee’s

Vision (some bees have been re-

ported to perceive ultraviolet but

not red) and they produce pleas-

ant, fresh fragrances. If néctar is

available it is usually produced in

a néctar tube at the base of the se-

pals or petals, though occasionally

as in Epidendram and its allies the

néctar tube is associated with the

ovary. Later we will discuss the

curious situation in many orchids

which do not produce néctar but

deceive the pollinator.

A considerable number of tropic-

al American orchids have adapt-

ed to peculiar behavior on the part

of male bees of the tribe Euglos-

sini. These will be discussed below.

Wasps — There are few orchids

exclusively adapted to wasps as

pollinators. Listera ovata, Coelo-

glossum viride, Epipactis latifolia,

and Herminium mcnorchis, Euro-

pean terrestrial orchids, have been

reported (Sprengel, 1793; Godfe-

ry, 1921; Darwin, 1862) to be pol-

linated by wasps. Wasps are at-

tracted to Ophrys by mimicry and

attempt to copulate with the flow-

ers (Godfrey, 1922; Kullenberg,

1961). Four Australian species of

Cryptostylis, called the tongue-or-

chid, are pollinated by the ichneu-

monid-wasp, Lissopimpla puncta-

ta (Coleman, 1927-1930). These

wasps attempt to copulate with the

flowers and thereby pollinate

them. Another Australian orchid

Calochilus campestris is pollinated

by the scoliidwasp Campsomeris

tasmaniensis.

I have recently observed taran-

tula hawks (wasps of the genus

Campsomeris — fig 1) as regular

visitors and pollinators of Brassia

cf antherotes in Colombia. Other

insects did not visit the flowers.

The wasps carried pollinia on the

front of their heads and deposited

them on the stigmas of other

flowers. The wasps do not hover in

front of the inflorescence but land

on the flower and then crawl from

flower to flower. The extraordinari-

ly long sepals of most brassias may

serve either to help or hinder the

progress of the pollinator. The

plants proved to have strong self-

incompatibility barriers with all

seed pods produced from flowers

pollinated with pollinia from the

same plant dropping off after

about 10 days. The long sepals may

tend to force the pollinator to fly

and thereby cause it to fly on to

other inflorescences. The same

wasp also pollinated Encyclia

pentotis.

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10 11 12 13 14 15

Volume 5 (Zoologia)

Fig. 1 — Campsomeris colombo, a wasp, pollinating flowers of Brassia a//.

antherotes in Colombia.

Dressler (personal communica-

tion) found wasps of Pachymerus

massadens visiting and pollinating

a species of Leochilus in Panama.

Wasps tend to be somewhat er-

ratic pollinators and are not as

easily guided as are bees (except

in cases of pseudocopulation) .

They are often seen collecting

néctar on the backsides of the se-

pals of orchids in the wild. They do

not, however, act as effective polli-

nators of these orchids. Van der

Pijl (1954) suggested that extra-

floral nectaries attract ants and

wasps which act to scare away

bees which would rob the flowers

by cutting into the nectaries from

the back of the flowers. I would

extend that hypothesis to include

the ants and wasps as guaráians

that keep away grasshoppers, cric-

kets, caterpillars and other insects

which eat the flowers. The sym-

biotic relationship between ants

and orchids which grow only in

ant nests, e.g. Coryanthes and

some species of Gcngora, Schom-

burgkia, and Epidendrum, may

depend on this defensive mechan-

ism. The flowers of these orchids

semm extremely susceptible to

damage by chewing insects when

the ants have been destroyed. Ants

have in no authenticated case prov-

en to pollinators of orchids.

Solitary bees — Many of the Eu-

ropean terrestrial orchids are pol-

em

6

Atas do Simpósio sobre a Biota Amazônica

linated by the solitary bees (bees

which do not have a social or col-

lective nesting systexn). Numer-

ous citations of pollination of Eu-

ropean orchids by solitary bees

have been published (see the list

of orchids and their pollinators) .

Members of the genus Cypripe-

dium have been reported to be pol-

linated by several species of An-

drena and Halictus. The bees are

attracted by deceit and after fal-

ling into the pouch, are forced to

crawl out under the stigma and

anthers. I found Phragmopedium

longifolium var. hartwegii to be

pollinated by halictine bees of the

genera Chlerogella and Chaenoha-

lictus.

Cálopogon parviflorus was re-

ported by Robertson (1887) to be

effectively pollinated by two spe-

cies of Auglochlora.

Diuris pedunculata of Australia

is pollinated by Halictus languino-

sus and Paracolletes sp. according

to Coleman (1932). She also ob-

served that only male bees were

attracted and were stupified while

on the flower.

I found Ptiloglossa ducalis, a

bee of the primitive family Colleti-

dae which flies at night and at

dawn, pollinating Miltonia endresii

Fig. 2 — Ptiloglossa ducalis, a night-flying colletid bee, pollinating Miltonia

endresii at 4 am in Costa Rica.

at about 4 am. in Costa Rica

(fig. 2). Other species of Miltonia

of the “Colombia” group are prob-

ably pollinated by night-flying

bees as well.

The pollination of several species

of Oncidium (O. planilabre, O. hy-

phaematicum and O. stipitatum)

by male bees of the genus Centris

is based upon the similation by the

flower of an enemy insect to be

driven away by male bees which

have established and are defending

a territory (Dodson & Frymire,

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Volume 5 (Zoologia)

7

1961 a and Dressler, personal com-

munication). I saw female bees of

the same genus, however, pollinat-

ing other species of Oncidium

such as O. ochmatochüum and

O. lanceanum as well as Odonto-

glossum grande in Costa Rica

(fig. 3). Centris is specially adapt-

ed to collect néctar from nectaries

iocated on the sepals of malpighia-

ceous flowers. This requires that

the bees land on top of the flower,

clutch the dorsal, stalked petal in

their mandibles, thrust their hairy

legs between the petals and strip

the néctar from the nectaries

Fig. 3 — The solitary bee Centris oncidophila pollinating Odontoglossum grande

in Costa Rica.

(fig. 4). Many Oncidium flowers

are strikingly similar to those of

malpighias — to the degree that

orchid collectors are often fooled

at a distance. The female bees land

on the Oncidium flower with the

callus holding them in position as

do the stigma and stamens of

the malpighiaceous flowers. They

clutch the column at the infra-

stigmatic lobe with their mandi-

bles and attempt to thrust their

legs between the petals and lobes

of the lip. Some oncidiums, e.g., the

O. ornithorynchum complex, have

developed nectaries between the

lobes of the lip and the column as

an apparant adaption to the habits

of the female Centris.

Semi-social bees — Carpenter

bees (the genus Xylocopa) occur

throughout the tropics of the world

Atas do Simpósio sobre a Biota Amazônica

and extend into temperate zones

in some areas. They drill holes in

timber to build their nests and are

often very large bees. They pol-

linate a number of the large-flow-

ered orchids of Asia such as, Eulo-

phia horsfallii (Kullenberg, 1961) ,

Vanda teres, Phajus tankervilliae,

Arundina speciosa, Phalaenopsis

amábilis (Van der Pijl, unpubl.),

Bromheadia spp. and Cymbidium

aloifolium (Ridley 1890 & 1896).

In tropical America they pollin-

ate Cattleyas, Sobralias, Epi-

dendrums, Encyclias and Schom-

burkgias (Dodson & Frymire,

1961a; Dodson 1965, unpubl.).

David Bennett (personal commu-

Fig. 4 — Centris versicolor visiting Malpighia glábra. The bee is collecting néctar

on its hairy legs by pulling them through the sepaline nectaries located between

the claws of the petals of the flower. The bees are often deceived by the flowers

of species of Oncidium which mimic Malpighia.

nication) collected Xylocopa tri-

cuspidifera pollinating Bletia cate-

nulata in Peru. In most cases xylo-

copas work large, strong orchid

flowers which produce néctar but

in a few cases, as in Oncidium

onustum and Barkeria lindleyana

(fig. 5) I found them to be attract-

ed more delicate flowers by sweet

fragrances. ( Xylocopa micans is

attracted to the pseudo-stamens on

the erect lip of Calopogon pulchel-

lus in Florida and while attempt-

ing to collect pollen are dropped

onto the column by their own

weight1 on the hinged lip. Probably

a majority of the large flowered,

bee-pollinated orchids of Asia are

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Volume 5 (Zoologia)

9

adapted to xylocopas and these

bees are also effective pollinators

of many similar flowers in the

Western Hemisphere.

The group of closely allied eu-

glossine bees Euglossa, Eulaema

and Euplusia is found only in the

tropical regions of the Western He-

misphere. All of the members have

extremely long tongues (more

than twice the length of the body

in some cases) and vary in size

frcm small to very large bees. The

male bees are peculiar in that they

are attracted to orchids, anthu-

riums, certain legumes and myr-

Fig. 5 — The carpenter bee Xylocopa micans visiting Barkeria lindleyana in

Costa Rica.

tles and even rotting wood or

bark, from which they get no di-

rect food. Of course, they also visit

néctar producing flowers for their

general sustenance. When male

bees visit non-nectar producing

flowers they scratch the surface of

the lip and sop up the exuded li-

quids with special pads located on

their front feet. The pads and the

swollen tibiae of the rear legs are

typical of male bees of the group.

After a very short period of such

behavior the male bees become in-

toxicated and loose motor control

and wariness.

Many genera of orchids such as

Stanhopea, Gongora, Coryanthes

(fig. 6) , Catasetum and Cycnoches

have adapted to this behavior by

developing complex pollination me-

chanisms. Normal bees would be

10

Atas do Simpósio sôbre a Biota Amazônica

too wary tc fali, slide or tumble

into and through the flowers, and

the systems are only effective after

the bees have been drugged. The

bees apparently enjoy the sensa-

tion for they constantly return to

the same flowers. This might easily

lead to self-pollination and there-

fore has resulted in further adapta-

tions, such as the separation of

sexes into different flowers in Ca-

tasetum and Cycnoches, and the

narrow stigma aperture which can

only be penetrated by thoroughly

dry pollinia of Stanhopea, Gongora

and allies.

Fig. 6 — Male Euglossine bees of Euplusia superba visiting Coryantes rodriguezii.

The bees scratch at the upper portion of the lip of the flower.

A eonsiderable list of orchids

pollinated by male bees of these

genera has now been compiled (see

List) . In all cases where the orchids

are adapted directly to euglossine

bees no other bees have been

observed visiting the flowers. In

most cases only a single species is

attracted to the flower and pollin-

ates it. In some cases several

species of bee may be attracted to

a flower but only one is of the

proper size to effect pollination. In

a very few cases, such as in Cata-

setum platyglossum (Fig. 7 — 8) ,

Gongora maculata (Fig. 9) and

Cceleopsis hyacinthosma (Fig. 10)

several species of euglossine bees

are attracted and are effective pol-

linators.

Euglossine bees of both sexes

visit rather unspecialized orchid

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Fig. 8 — A male bee of Eulaema cingulata scratching inside the hood-shaped lip

of the female flower of Catasetum platyglossum. Note the pollinarium írom a

male flower attached to the thorax of the bee.

Fig. 7 — A male Euglossine bee, Eulaema cingulata, scratching at the lip of the

male flower of Catasetum platyglossum. The bee has touched the antennae which

discharges the pollinarium. The sticky viscidium of the pollinarium has just

struck the bee and will become glued to the thorax of the bee.

cm

12 Atas do Simpósio sôbre a Biota Amazônica

Fig. 10 — A male Eulaema meriana scratching at the lip of Coeliopsis hyacin-

thosma in Panama.

Fig. 9 — Male Euglossa decora visiting Gongora quinquenervis and scratching at

the inside of the lip.

cm

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13

Fig. 11 — Male Euplusia surinamensis entering the flower of Sobralid : Violacea.

Note the pollinarium attached to the thorax of the bee.

Fig. 12 — Male Eulaema polychroma with pollinaria on the thorax entering the

flower of Cattleya loarszewiczii.

14 Atas do Simpósio sôbre a Biota Amazônica

flowers such as those of the genera

Sobralia, (Fig 11) Bletia and Cat-

tleya (Fig. 12) which produce

néctar and are visited by other

bees as well. Both sexes also visit

more specialized néctar producing

orchids, such as Aspasia and Tri-

chccentrum (Fig. 13). They have

been observed pollinating several

species of Maxillaria, Lycaste, An-

guloa (Fig. 14) Zygopetalum and

related genera, (Fig. 15) as well as

Trichopilia, Notylia (Fig. 16 -j- 17)

and Lockhartia, none of which

produce néctar but tend to in-

toxicate the male bees. Catase-

tum, (Fig 7 + 8) , Cycnoches,

(Fig. 18 + 19) , Mormodes (Fig. 20)

and all of the Stanhopea alliance

are exclusively pollinated by eu-

glossine bees and have developed

some of the most remarkable pol-

lination mechanisms to be found

in the plant kingdom is response

to the behavior of the bees.

Social Bees — Social bees are

among the most important pollin-

ators of flowers in general but

peculiarly have not achieved the

same prominence in regard to pol-

lination of orchids. Only a few

records of orchids pollinated by so-

cial bees are available. It may be

that social bees tend to be too

constant to néctar producing flow-

ers and therefore have tended to

ignore poor providers such as or-

chids.

Fig. 13 — Male Eulaema cingulata visiting the flower of Trichocentrum tigrinum

in Ecuador.

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Volume 5 (Zoologia)

15

Pig. 14 — Male Eulaema boliviensis scratching at the inside of the ílexible lip of

Anguloa clowesii in Colombia.

Fig. 15 — Male Eulaema polychroma pollinating a flower of Pescatorea wallisn.

16

Atas do Simpósio sôbre a Biota Amazônica

Fig. 16 — Male Euglossa augaspis hovering before flowers of Notylia buchtenii.

Note the pollinarium attached to the head of the bee.

The common honeybee, Apis me-

lifera, is now distributed through-

out the tropical and temperate

zones of the world. Its origin, how-

ever, was probably in the Near

East or Asia. Consequently orchids

in the New World are not adapted

to it. In Asia Apis dorsata pollin-

ates Dendróbium crumenatum

and Cymbidium finlaysonianum

(Burkhill, 1919) . Apis dorsata and

A. indica were observed by Burk-

hill to pollinate Dendróbium su-

perbum. Ridley (1905) reported

A. dorsata as the effective pollina-

tor of Grammatophyllum specio-

sum. Honeybees are often attract-

ed to the pleasant odors of tropical

orchids in the green house but

have not been reported as pollina-

tors of these orchids in nature.

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Fig. 17 — Male Euplusia surinamensis pollinating flowers of Notylia xytriophora

in Ecuador.

Fig. 18 — a: Male bee of Eulaema cingulata on male flower of Cycnoches

lehmanii. b: The same bee with pollinarium attached to the abdómen about

to pollinate the female flower of Cycnoches lehmanii.

The bumblebees ( Bombus ) are

also distributed throughout the

temperate and tropical zones of

the world. In the temperate zones

they have been reported as the pol-

linators of several species of Orchis

and Spiranthes. They also pollinate

Epipogium aphyllum (Knuth, . . .

2 — 37 145

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Fig 19 — a: Male bee of Eulaema nigrita visiting the male flowers of Cycnoches

aureum. b: The same species of bee visiting female flowers of the same

Cycnoches.

Fig. 20 — Male bee of Euglossa hemichlora scratching at the lip of Mormodes cf.

buccinator in Ecuador. Note the freshly attached and curled pollinarium on the

thorax of the bee.

1898) and Goodyera repens (Dar-

win, 1862). In tropical America

they are effective pollinators of

such generalized orchids as Sobra-

lia along with other large bees

(Dodson, 1962 a & 1965) . I observ-

ed Bombus hortulans var. robustus

as the pollinator of Odontoglossum

cm l

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Volume 5 (Zoologia)

19

kegeljani (Fig. 21). I also found

that the same species acts as a

co-pollinator with Centris of On-

cidium macranthum and that

Bombus volucellioides pollinates

Maxillaria fletcheriana in the An-

des of Ecuador. Bombus will prob-

ably prove to be an important

pollinator of large-flowered, non-

-specialized orchids at higher eleva-

tions in tropical America. It also

visits and pollinates large-flow-

ered orchids in green houses in

temperate zones.

Trigonas are relatively small,

stingless bees of tropical America.

They tend to be rathcr erratic

fliers but are often quite specific

as to the flowers they visit. I en-

countered Trigona testacea and

T. amalthea as pollinators of Ma-

xillaria reichenheimiana in Ecua-

dor in 1963. Trigona cf. trinidaden-

sis effectively and exclusively pol-

linates Xylobium latilabium (Fig.

22) in Peru even though there are

several other species of Trigona

nearby at all times which are not

attracted to it.

I observed a male bee of Trigona

testacea visiting flowers of Coelo-

gyne lawrenciana (a native of

Asia) in Costa Rica in 1963. The

bee was in a highly excited state

and entered flower after flower

only to leave each moments after

entering. The bee functioned as a

very effective pollinator.

Kerr & Lopez (1963) reported

Trigona doryana (males only) as

Fig 21 — The bumblebee (social) Bombus hortulans var. robusta pollinating

Odontoglosswm kegeljani.

20

Atas do Simpósio sôbre a Biota Amazônica

Fig. 22 — Trigona amalthea, a tropical social bee, pollinating Xylobium

latüabium in Peru.

the pollinator of Trigonidium obtu-

sum. The bees attempted to copu-

late with both paired petals, which

are glandular and produce odor.

Meliponas are médium size,

stingless social bees of tropical

America. One common species, M.

beecheii has been supposed to be

the natural pollinator of Vanilla in

México. Melipona eburnea exclusi-

vely pollinated Maxillaria fuersten-

bergae (Fig. 23) in Ecuador (Dod-

son 1962 a). Melipona flavipennis

pollinates Cattleya luteola at dawn

in Peru (Fig. 24). Meliponas were

reported by Ducke on diverse

nectar-secreting orchids in South

America. The genus may be impor-

tant as pollinators of maxillarias.

MOTHS AND BUTTERFLIES

Moths and butterflies are closely

related and are not always clearly

distinct, day-flying hawk-moths

and skippers tending to bridge the

differences. Nevertheless, their be-

havior tends to be so distinctive

that the flowers adapted to them

are easily separated into two clas-

ses. Both moths and butterflies

have tubular tongues or probosci

which are curled when not in use.

When extended into the néctar

tube of a flower the probosci func-

tion as tubes through which the

insects suck up the néctar.

Moth and butterfly flowers —

Moths normally fly and feed at

night and are attracted to flowers

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Fig 23 — Melipona eburnea, a social bee, pollinating Maxillaria furstenbergae.

that produce strong odors and are

white or light colored. Moths that

visit flowers are usually strong

fliers capable of hovering in front

of the flower while extracting the

néctar. The typical moth-flower

has a long slender néctar tube

containing abundant néctar. The

fragrance produced is typically

sweet or musky and the flowers are

usually horizontal or hanging. But-

terflies, on the other hand, are day

fliers and go to highly colored

flowers which may or may not be

fragrant. They tend to be some-

what erratic fliers and usually land

on the flowers and therefore the

flowers are usually erect and pro-

vide platforms for landing. Often

the platforms simply consist of a

head of erect, tightly packed

flowers. Butterflies detect colors

well and flowers adapted to them

for pollination are usually brightly

24 Atas do Simpósio sôbre a Biota Amazônica

Fig. 25 — The butterfly Papilio polyxenes var. americus pollinating Epidendrum

secundum in Ecuador.

Bird-pollinated flowers — The

birds that act as effective pollin-

ators of flowers usually have long

slender beaks that are introduced

into the flower. Hummingbirds

hover in front of the flowers while

other birds land on the flowers or

on nearby stems. The flowers

adapted to other flower-visiting

birds either provide a special land-

ing platform (i.e., Strelitzia) or are

produced close to the stems of the

plant or other substrates. Birds are

not known to have a sense of smell

but have excellent vision and flow-

ers adapted to them are usually

very brightly colored with red, blue

and yellow predominating. Bird-

-flowers are usually tubular in

form, often with exserted anthers

that brush pollen on the head or

body of the bird. The styles of these

flowers are likewise exserted.

Bird-pollinated orchids tend t o

follow the pattern of other bird-

-flowers. In some cases, however,

they diverge considerably. Many

orchids of the Western Hemisphere

appear to have adapted to bird-pol-

lination as an extention of butter-

fly-pollination and as in the case

E. secundum birds and butterflies

act as co-pollinators. In such cases

orchid flowers already adapted to

butterflies are not greatly changed

morphologically. On the other

hand, orchids which have adapted

directly to hummingbirds from

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bee-pollinated ancestors change

fundamentally (Dodson, 1962 a) .

Cochlioda, Sophronitis, L a eli a

(Section Cyrtolaelia) , Elleanthus

(Pig. 26), Isochilus, certain species

of Masdevallia , Rodriguezia, Sobra-

lia and Maxillaria and several

small genera such as Comparettia,

Hexisea, Alemania, Meiracilium

and Nageliella are all remarkably

similar in certain aspects. All have

bright colors, tubular flowers and

a callus or hump in the interior of

the tube on the lip that acts to

force the beak of the bird against

the column (Fig. 39). The callus

or hump and bright colors are also

present in bird-pollinated orchids

from the Old World tropics.

Hummingbirds — I have made a

number of observations of orchids

Fig. 26 — Elleanthus capitatus, a hummingbird pollinated orchid typical of other

members of the genus in flower structure.

being pollinated by hummingbirds.

Eight species of Elleanthus were

observed being pollinated by hum-

mingbirds and a certain constancy

on the part of the birds to colors

in sympatric species was noted.

Masdevallia rosea and Epidendrum

secundum are pollinated by hum-

mingbirds in Ecuador (Dodson,

(1962 a) and Cochlioda vulcanica,

Rodriguezia secunda, Isochilus car-

nosifolius, Maxillaria sp., Compa-

rettia falcata, (Fig. 27) and Sobra-

lia amabilis were observed being

pollinated by hummingbirds in

Ecuador, Costa Rica or Peru.

26

Atas do Simpósio sobre a Biota Amazônica

Sugarbirds — These birds were

considered to form the family Coe-

rebidae of the New World tropics.

Recently, however, this family has

been broken up and several species

placed with the Tanagers. The

birds visit flowers for néctar and

in one instance I observed them

visiting Elleanthus roseus in Ecua-

dor.

Sunbirds — These birds belong

to family Nectarinidae and occur

in África and tropical Asia. Probab-

ly a considerably number of or-

Fig. 27 — Comparettia falcata, an orchid pollinated by the hummingbird

Ama.za.lia tzacatl. Note the concave callus on the lip under the column which

forces the beak of the bird against the collumn.

chids are adapted to them but only

the reports of Cyrtostomus pecto-

ralis visiting Dendrobium secun-

dam by Burkhill (1919) and of

sunbirds visiting Dendrobium has-

seltii in Java (Docters van Leewen

1933) are available. Dendrobium

lawesii was reported by Slade ....

(1963) to be bird-pollinated at high

elevations in New Guinea but the

bird was not identified and may

belong to this family or the fol-

lowing.

Nectar-eaters — These birds be-

long to the family Meliphagidae

and occur in the Pacific Islands.

Unless the bird mentioned above

in the report by Slade belongs to

this family no reports of orchids

being pollinated by nectar-eaters

are available. Numerous Dendro-

biums, Cryptochilus and other or-

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27

chids with flowers of construction

and color similar to known bird-

-pollinated orchids occur in their

range and may well be visited by

them.

Honeycreepers — These birds

belong to the family Drepanididae

and are found in Hawaii. They

have not been reported pollinating

orchids and none of the Hawaiian

orchids have the birds-pollination

syndrome.

FLIES

Some flies are important pollin-

ators of flowers and certain fami-

lies such as the Syrphidae and

Bombylidae are restricted to flow-

ers for their food.

Fly-pollinated flowers — Fly-

-pollinated flowers in general are

called myophilous. Unspecialized

flowers often attract flies, bees,

beetles and butterflies. These are

usually shallow, produce abundant

néctar on surface nectaries and

emit sweet odors. The flies eat the

néctar and do not store it as do

bees. More specialized “fly-flowers”

may attract flies through decep-

tion, imitating decaying substan-

ces, dung or carrion. Many kinds of

flies, midges, gnats, mosquitos,

etc., are then attracted and act as

effective pollinators. These flowers

are called sapromyophilous and are

most numerous in warm, arid or

tropical zones, especially in África.

Sapromyophilous flowers often

have developed traps for catching

and holding unadapted visitors or

are radial in form. They commonly

have large landing surfaces and

“tails” produced from the flower

parts which function as guides.

Their colors are usually dull green,

brown, purple or red with check-

ered or blotched designs. The odors

produced are commonly putres-

cent. Flies, unlike bees, are some-

what erratic fliers and are poorly

guided.

Orchids pollinated by flies are

common throughout the world. In

most fly-pollinated orchids special

adaptations have developed — su-

perimposed on the basic bee-pollin-

ated orchid flower — to guide po-

orly oriented flies. Certain of the

five tepals may be long and “tail”

like ( Bulbophyllum and Masdeva-

lia) or joined to form a flat, radial

flower ( Stelis ). The flowers them-

selves may be arranged to form a

radial, compound “flower” ( Cir -

rhopetalum ) . The petals or lip may

be fringed with motile club like

hairs that vibrate in the wind and

attract the flies ( Bulbophyllum

and Cirrhopetalum ) . Often the

sepals are joined or the lip is sac-

cate to form a trap ( Phragmope -

dium and Pterostylis) into which

the flies fali and must crawl out

by means of a tunnel that passes

the stigma and anther. A common

contrivance is a hinged, balanced

lip which tips with the weight and

28

Atas do Simpósio sôbre a Biota Amazônica

launches the pollinator into the

flower.

In orchids it is often difficult

to decide where simple myophily

ends and sapromyophily begins.

Many genera have some species

whose pollination is based on

simple myophily, attracting va-

rious flies on the basis of sweet

odors and néctar production, while

others attract flies on the basis of

rotten odors and provide no food.

Three major groups of orchids

have become predominantly myo-

philous i.e., the Pleurothallidinae

(in America), the Bulbophyllum

group (mainly in the Old World)

and the large genus Pterostylis and

its relatives (in Australia). Other

genera have only a few species

adapted to flies while the majority

of species may be adapted to other

pollinators.

I found Phragmopedium longifo-

lium var. hartivegii to be pollinat-

ed by flies of the genus Syrphus

but it was also pollinated by bees.

The flies and bees, apparently

searching for food, slipped into the

saccate lip and in crawling out

deposited pollinia on the stigma

and piked up a new set.

Two species of the genus Bnlbo-

phyllum, B. macranthum and B.

emiliorum, have been reported to

be pollinated by flies (Pohl, 1935;

Ridley, 1890 b) . In both cases the

flies were thrown against the

ventral surface of the column when

the hinged lip was overbalanced.

I described similar phenomena in a

greenhouse grown plant of Cirrho-

petalum ornatissimum visited by

common house flies (Fig. 28). It

is presumed that most of the more

than 1,000 species in this group are

pollinated in a similar manner.

The Pleurothallidinae also con-

tain nearly 1,000 species of fly-pol-

linated orchids. Three species of

Pleurothallis, P. xanthochlora, P.

ruscifolia and P. eumecocaulon,

have been reported to be pollin-

ated by Drosophilas (Dodson, ....

1962 a & 1965) and Pleurothallis

monocardia (Fig. 29) is pollinated

by Lyccria sp. In the same group

of orchids Masdevalia fractiflexa

and M. erythrochaete are pollin-

ated by flies, the former attracting

carrion flies to its foul odor and the

latter Drosophilas to its basket-

-like, néctar producing lip.

Members of the genus Pterostylis

of Australia have developed a

complex trap composed of joined

sepals. Pterostylis mutica was re-

ported by Hyett (1960) to be pol-

linated by flies of the families Psy-

chodidae and Mycetophilidae. Sar-

GENT (1909), COLEMAN (1934 b)

and Werth (1951 & 1956) also

described fly-pollination of mem-

bers of this genus. The related

genus Corybas was reported to be

pollinated by gnats (Family Culi-

cidae) by Thompson (1927).

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Garnet (1940) reported the pol-

lination of four species of Praso-

phyllum in Australia by flies of the

family Chloropidae. In this genus

the lip is uppermost in the flower

and is flexible. When the fly lands

on the lip it deflexes dropping the

fly on the column where the pol-

linia are deposited. Ridley (1894)

described the pollination mecha-

nism of two Malayan species of

Plocoglottis, i.e.} P. foetiãa and P.

porphyrophylla, in which the lips

are under tension between the pe-

tals. When the fly lands on the lip

its weight releases the connections

and the lip springs upward, thrust-

ing the fly against the column.

Several reports of species from

genera otherwise known for bee or

Lepidoptera pollinated flowers

Fig. 28 — Bulbophyllum (.Cirrhopetalum) ornatissknum being pollinated by flies.

Note the flies trapped in the flowers by the over-balanced lip of the flower.

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30

Atas do Simpósio sobre a Biota Amazônica

have been made. Epidendrum fim-

briatum is pollinated by flies of the

Fig. 29 — A Sciarid fly, Lycoria sp.

pollinating the flowers of Pleurothallis

monocardia. Note the polinia attached

to the legs of the fly.

family Tachinidae (Dodson, 1962a)

and Vogel (1954) reported Maxil-

laria lepidota and Disa lugens to

be fly-pollinated. Gastrodia javani-

ca is fly-pollinated according to

Holtum (1953) while Dendrobium

linguiforme is pollinated by Syr-

phus viridiceps according to Gil-

bert (1958). Habenaria obtusata

is pollinated by mosquitos in Mi-

chigan (Dexter, 1913).

Trichoceros antennifera mimics

the female flies of Paragymnomrp,a

sp. of the family Tachinidae and

the male flies attempt to copulate

with flowers, thereby pollinating

them quite effectively (Dodson,

1962 a).

BEETLES

Beetle-pollinated flowers are

called cantharophilous and tend to

occur in the more primitive fami-

lies of flowering plants. Coleman

(1933) reported the beetles Ameta-

lia and Trogoderma as pollinators

of Prasophyllum odoratum in Aus-

trália. Darwin (1877) reported a

beetle, Strangalia atrata as a co-

pollinator of Orchis mascula. Rid-

Ley (1894) observed beetles (Rhy-

nocophorus sp.) creeping about

among the nectarless flowers of

Dendrochilum longtfolium with

pollen masses on their heads and

Bolus (1839) found a beetle on

Disa bivalvata in South África.

These are the only reports of

beetle-pollination to date for the

orchids. The cases are obviously

scattered and there appears to be

no trend toward adaptation to be-

etles as pollinators.

OTHER POLLINATORS

Reports of orchid pollination by

snails and ants seem to have no

foundation in fact. A recently pu-

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blished report of frog-pollination

of Lissochilus roseus seems incredi-

ble at best and may have been a

simple misinterpretation of a phe-

nomena reported by Jones (writ-

ten communication) where lizards

were observed on numerous occa-

sions on the inflorescences of

Schomburgkia in the Barbados

Islands. The lizards were consu-

ming ants which were attracted to

the extra-floral nectaries on the

outside of the sepals and bracts.

The possibility of such animais

acting as pollinators in the normal

sense seems remote. I have often

seen spiders waiting for prey inside

the open flowers of Sobralia and

Cattleya but they do not function

as pollinators.

There have been no reports of

orchid pollination by bats, wind or

water. These pollinating agents

would not be expected for the flor-

al type of the orchid would not

readily lend to adaptation to them.

AUTOGAMY

Self-pollination occurs in a sig-

nificant number of orchids. Several

degrees of this phenomenon may

be found in a single genus, from

species in which accidental self-

-pollination results in fertilization

to those in which the flowers never

open yet produce fertile seed. In

many orchids self-fertilization pre-

sumably is not possible due to gen-

etically controlled self-incompat-

ibility in which pollen from a plant

with a particular combination of

genetic factors will not fertilize its

own ovules or those any other

plant with the same combination.

In most species the pollen is kept

separate from the stigma by the

rostellar flap. This physical barrier

is normally quite effective but in

some species forms occur in which

the rostellum degenerates or be-

comes stigmatic and self-pollina-

tion results when the pollen germi-

nates on contact with the stigma-

tic fluid. In most of these forms

normal plants are found in the

same population. Self-pollination

may also occur as a result of simple

falling of old disintegrated pollinia,

a means of averting sterility at the

end of a long, normal period when

the flower is open and no pollin-

ator arrives.

A kind of mechanical self-pollin-

ation occurs in some orchids where

the caudicles connecting the visi-

dium and the pollinia bend down-

ward and carry the pollinia into

place on the stigma. This has been

observed in Ophrys apifera (van

der PiJL, unpubl.) Herminium,

Epipactis, Orchis (Martens, 1926) ,

Oncidium glossomystax (Dodson,

unpubl.), and Oncidium ostlun-

dianum (Dressler & Wirth, per-

sonal communication).

Generally speaking, autogamy

seems a means of averting extinc-

tion in plants growing under con-

32

Atas do Simpósio sobre a Biota Amazônica

ditions adverse for normal pollin-

ation relations (Stebbins, 1950

and Grant, 1963). Examples are

some species of Orchis in Europe

and several orchids in Florida,

such as Epiãendrum nocturnum,

Encylia cochleata and Bletia pur-

púrea, all of which have non-au-

togamous forms in other areas.

Seed blows in to Florida from the

Caribbean region where normal

pollinators exist, and those plants

that survive the lack of pollinators

in Florida tend to be autogamous.

Probably more than 200 species

of orchids have been reported to

more or less regularly autogamous.

Undoubtedly many more exist.

Many of these forms of normal

species — with their characteristic

changes in morphological struc-

ture — have been given names and

are frequently regarded as distinc-

tive species.

A POLLINATION SPECTRUM

The published data are insuf-

ficient to establish accurately the

relative importance of pollinating

agents throughout the family.

However, by surveying the syn-

drones of the flowers of the family

in general, it is possible to give a

rough estimate in percentages of

the kinds of pollinating agents to

which the orchids are adapted.

This is called a “pollination spec-

trum”.

Hymenoptera

Wasps 3%

Solitary bees 10%

Carpenter bees 12%

Euglossini 5%

Social bees 13%

Mixed bees 11%

54%

Other Agents

Moths 15%

Butterflies 3 %

Birds 3 %

Flies 12%

Mixed agents 11%

46%

Such extreme divergence in pol-

linators is quite uncommon in

other families of flowering plants

and testifies for relative youth,

continuing evolution and adaptive

radiation in the family. The pol-

lination-spectrum of the family

Polemoniaceae (Grant, 1961) is

based on more thorough and con-

centrated study of a much smaller

and in some respects diverse group.

ATTRACTION OF THE

POLLINATOR

Most of the pollinators of the or-

chids are attracted to the flowers

on the basis of the same instincts

that attract them to other flow-

ers. Food-gathering is their pri-

mary interest. Visual signals pro-

vided by the flowers, reinforced by

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characteristic fragrances, act on

instinctive backgrounds as the

physical attractants. In some cases

food is available and in others it

is not. In some orchids, the re-

productive phase of the life-cycle

of the pollinator is interrupted and

orchid flower plays upon those

instinctive behavior patterns which

guide male insects during mating.

VISUAL ATTRACTION

Visual stimuli are usually asso-

ciated with fragrances but not al-

ways. Sensitivity for certain colors

is important in the attraction of

pollinators with differences in per-

ception of the color spectrum play-

ing a crucial role. Bees, for exam-

ple, apparently do not perceve pure

red but do see into the ultraviolet

end of the spectrum. Butterflies

and birds, on the other hand, are

strongly attracted to red and bril-

liantly colored orchids can gener-

ally be expected to be pollinated

by those agents. Flies are attract-

ed to dull-red, green, and purple,

but here color is often reenforced

by motion in the form of streamers,

hairs, articulated b r a c t s, etc.,

which flutter in the breeze. It can

be said that visual stimuli play an

important role in the selective

attraction of major groups of pol-

linators. I have gathered evidence

that color may act as a barrier to

hybridization by attracting certain

species of humming birds and not

others. Two species of Elleanthus;

E. roseus and E. arapophyllosta-

chys, are sympatric on the western

slopes of Mt. Pichincha in Ecuador

and flower at the same time. The

plants are often intermingled.

Though the flowers are morpholo-

gically very similar they differ in

color, the former being pink and

the latter orange. The former was

regularly visited by an unidentified

hummingbird and the latter by

Ocreatus undervooodii. No hybrids

were found and the birds were not

observed to visit the opposite

species.

In cases where pollinators are

attracted to orchid flowers on the

basis of sexual instincts the flow-

ers will usually exhibit the essen-

tial visual characters that attract

the male of the species to the fe-

male ready for mating. Kullen-

berg’s (1961) work on pseudocopu-

lation in Ophrys indicated that

specific color patterns in the ultra-

violet end of the spectrum played

a role in attraction of specific male

bees and wasps. Further stimuli in

the form of vestiture on the lip of

the flower upon which the insects

rub and odor also form part of the

overall attraction spectrum and

are necessary for culmination of

the behavior leading to deposition

of the pollinia.

3 — 37 145

34

Atas do Simpósio sôbre a Biota Amazônica

ODOR

In orchids, fragrances often dom-

inate and may even cancel color

effects in the attraction of the pol-

linators. In some genera such as

Stanhopea, Gongora, Coryanthes

and Catasetum, red is a common

color but field experiments (Dod-

son & Frymire 1961 a) indicate

that the bees will search out the

flowers by following up the fra-

grances even if the flowers are

hidden. Consequently the presence

of red here seems not to be involv-

ed in visual attraction. Fragran-

ces produced by individual species

have apparently played a very im-

portant role in speciation in many

orchids. Where attraction on the

basis of odors is dominant the spe-

cies can easily be distinguished on

the basis of odors alone by the use

of gas-liquid - chromatography in

their analysis (Rasmussen & Dod-

son, unpubl. and Dodson unpubl.)

The odors produced are often very

complex and contain distinct Che-

mical components in varying

quantities. Some species of Stanho-

pea, for instance, were found to

have as many ten components in

their odor, while others had as few

as three. In each, different specific

pollinators are attracted. Some or-

chids such as certain forms of Gon-

gora maculata produce odors not

perceptible to humans yet attract

bees that are attracted normally

on the basis of odors.

Extreme cases of orchid species

which are not distinguishable

under normal taxonomic criteria,

yet are reproductively isolated

through the attraction of different

species of bees as pollinators, have

been encountered in Gongora and

Stanhopea (Dodson, 1965 a) and

Dressler, personal communica-

tion). After the pollinators have

demonstrated their preferences it

is possible to separate the taxa on

the basis of characters which have

been given minor importance as

criteria. Each of these cryptic spe-

cies has a distinctive odor. The

significance of these phenomena

in taxonomy will be discussed

below.

FOOD

Orchids produce néctar as food

for their pollinators as do other

flowering plants. Pollen, however,

is not made available and on the

contrary, is usually protected by

an anther. Pollinators that visit

orchids for food then must visit

other flowers in order to collect the

pollen which provides protein in

their diet (and in the case of bees,

in the diet of the larvae).

Néctar — Néctar, in the orchids,

is produced on the lip, in nectar-

-tubes (spurs) developed from the

back of the lip or a combination

of lip and sepals, or in néctar tubes

beside the ovary. It is important

to note that in the case of Epiãen-

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drum and its allies, the nectaries

are not within the septa of the

ovary itself as they are in many

other flowering plants. In some or-

chids, such as Deudrobium, néctar

is produced on an extention of the

column called the mentum, chin

or foot. Sometimes, as in species of

Bulbophyllum and Pleurothallis,

the néctar is produced on the sur-

face or in grooves or on the callus

on the lip, petals and sepals. In

other cases, as in some species of

Epidendrum, the néctar is not free

at all even when in néctar tubes

but is contained in soft-walled

cells which are broken by the

tongue or proboscis of the pollin-

ator.

Pollen-imitation and food hairs

— Several orchids do not produce

néctar but have a powdery mass

resembling pollen on the lips of the

flowers. Each grain is a soft-walled

cell containing starch and is ap-

parently collected by the bees.

Many species of Maxillaria (Porsch

1905) produce pseudo-pollen of

this type on the lip as do some

species of Polystachya and Eria

(Beck, 1914). In some cases the

cells are detached papillae and in

others they are disintegrated mul-

ticellular hairs.

Beck (1912) assumed that the

hairs of the lip of species of Vanilla,

Coelogyne and Cymbidium are

grazed on by bees. He found the

hairs to be thin-walled and filled

with plasma and sugar. No obser-

vation substantiating his assump-

tions have been made.

Callus — It has often been as-

sumed that the callus on the lip

of most orchids is there as food

for the pollinators. I have, on the

contrary, not found any instance

where the actual pollinators of an

orchid eat or gnaw on the callus.

In all cases observed to date the

callus has functioned in guidance

of the pollinator and in many cases

acts as a semi-barrier to make the

pollinator more effective. Reports

(e.g., Cruger, 1865) of the pollin-

ators “gnawing” the callus of the

orchid flower are probably attrib-

utable to faulty observation.

DECEPTION OF THE POLLINATOR

As mentioned above, the majori-

ty of the orchids produce food as

the primary attractant of the pol-

linator. In many orchids, however,

deception of the pollinator is the

primary means of attraction. Often

the pollinator must expect to en-

counter food where none exists. In

the less complex cases, odor, color

and form of the flower often are

similar to those of other flowering

plants nearby which do produce

food. This system has been remark-

ably successful in the family and

can be encountered throughout

their geographical distribution, but

is particularly obvious in the tro-

pics. Deception seems fundamental

36

Atas do Simpósio sobre a Biota Amazônica

to the development of many of

the more complex pollination me-

chanisms in the orchids.

Food-deception — In many or-

chids such as Cattleya no obvious

néctar exists though liquid is pre-

sent in thin-walled cells on the

walls of the nectar-tube. These

cells are presumably ruptured by

the tongue of the pollinator and

thereby release a small quantity

of néctar. This may be a step in

the direction of food-deception. In

the subtribe Oncidinae certain of

the less advanced members have

nectaries while the majority are

based on food-deception with some

species based on pseudoantagon-

ism. Remarkably, some advanced

species of Oncidium have reverted

to the production of néctar but

now produce in on the lateral lobes

of the lip. Food-deception is en-

countered in other plant families

such as the Euphorbiaceae, Ara-

ceae and Aristolochiaceae but is

uncommon when compared with

the situation in the orchids. It

seems a necessary phenomenon in

a series of steps leading to higher

forms of deception as mimicry of

females and antagonists. A clear

transition from mechanisms based

on simple food-deception to the

most complex of mechanisms based

on probable initial sexual attrac-

tion reinforced by intoxication

(i.e., Stanhopea and Catasetum) is

found in the subfamily Epiden-

drioidae.

In Calopogon, one of the prettier

temperate zone orchids of the east-

ern United States and Canada, the

lip is uppermost and has fleshy

protuberances projecting and co-

lored so as to appear as a mass of

stamens. Robertson (1887) report-

ed several bees that attempted to

land on the lip to collect pollen

from the false stamens of C. par-

vtflorum. I observed carpenter bees

attempting to collect pollen from

the false stamens of C. pulchellus

in Florida. The lip is articulated

at the base and the weight of the

bee causes it to depress, thereby

lowering the bee into the column.

In attempting to escape, the bee

deposits pollinia in the stigma and

picks up a new set on the upper

thorax.

A similar type of food-deception

occurs where flies are attracted to

flowers which smell like rotten

meat or other putrescent substan-

ces. The flies may or may not lay

eggs in the flowers. No actual food

exists, however.

Pseudocopulation — Pseudoco-

pulation is one of the more ad-

vanced forms of deception of the

pollinator. It has only been report-

ed extensively in the orchids and is

based upon mimicry of the female

insect. (The use of the term “mi-

micry” has been long surrounded

with controversy. In Zoology the

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Volume 5 (Zoologia)

37

term signifies a close externai re-

semblance of a animal to some

other animal or object serving for

protection or concealment as a re-

sult of selective adaptation. In

horticulture, unfortunately the

the term has been used to describe

the similarity of a flower or its

parts to a commonly known object,

such as, the callus of Peristeria

elata to a dove, the callus of Odon-

toglossum grande to a baby in a

cradle, the flower Coeloglossum vi-

ride to a frog, the flower of Pha-

laenopsis amabilis to a moth, etc.

These similarities are entirely for-

tuitous and cannot be considered

as significant in the biology of the

flowers involved. I use the term

here to refer to characteristics se-

lectively developed in a flower to

deceive the pollinators) . Pseudo-

copulation has been well described

in Ophrys (Correvon & Pouyanne,

1916; Godfrey, 1933 and Kullen-

berg, 1961), Cryptostylis (Cole-

man, 1927, 1928, 1929 a & b and

1930 a & b) , Diuris (Coleman,

1932) , Trichoceros (Dodson, 1962a)

and Trigonidium (Kerr & Lopez,

1963) .

The remarkable resemblance to

insects of the flowers of various

species of Ophrys is obvious by the

common names which have been

applied to them, i.e., Bee-orchids,

Fly-orchis, etc. In a letter cited by

Rolfe (1910) the belief was ex-

pressed that the likeness of the

flowers to insects has nothing to

do with pollination but served to

frighten away browsing cows. Dar-

win (1872) wrote in a footnote that

“Mr. Price has frequently wit-

nessed attacks made upon the bee-

orchis by a bee, similar to the trou-

blesome Apis muscorum”. Darwin

also noted that he could not con-

jecture what this meant.

Pouyanne discovered (Correvon

& Pouyanne, 1916) in Algeria that

flowers of Ophrys speculum, offer-

ing no food whatever, attract a

male wasp by imitating its females.

Later Godfrey & Wolff cor-

roborated the observations. Kul-

lenberg (1956 & 1961) studied the

problem from the zoological point

of view. He found that the Ophrys

flower attracts the male bee or

wasp by odors similar to those pro-

duced by the females. The odors

alone tend not to be species specific

and often attract several kinds of

bees. Shape, colors and vestiture

on the lip tend to complete the

attraction and exclude any other

than the correct pollinator. The

behavior of the pollinator is similar

to that during the initial phases of

copulation with females.

As the pollinators attempt to

copulate with the lip, the pollinia

are deposited on the stigma and

a new set is carried away. Four

genera of solitary bees and wasps

appear to be the principal pollin-

ators. Ophrys speculum lures the

38

Atas do Simpósio sôbre a Biota Amazônica

digger wasp Scolia cüiata (Kul-

lenberg, 1961). Ophrys muscifera

and O. insectifera are pollinated by

the wasp Gorytes mystaceous

(Godfrey, 1933) . Ophrys fuciflora,

O. tenthredinifera, O. bombyliflo-

ra, O. scolopax and O. apifera are

pollinated by bees of the genus En-

cera (Godfrey, 1933 and Kullen-

berg, 1961). Ophrys arachnitifor-

mis, O. lutea and O. fusca are pol-

linated by bees of the genus An-

drena (Kullenberg, 1961).

The species of Ophrys which are

pollinated by Scolia, Gorytes and

Encera induce the insects to at-

tempt copulation with the apex of

the lip and the pollinarium is de-

posited on the head of the insect.

Those pollinated by Andrena ap-

pear, for the most part, to stimu-

late the bee to reverse the position

and copulate with the base of the

lip and the pollinarium is attached

to the abdômen. Dimensions of the

flowers help in determining speci-

ficity and success.

Pollination of four species of the

genus Cryptostylis of Australia was

reported by Coleman (1927, 1928,

1929 a & b and 1930 a & b). She

observed males of Lissopimpla se-

mipunctata, an ichneumon wasp,

attempting to copulate with the

flowers of Cryptostylis leptochila,

C. subulata, C. erecta and C. ovata.

The wasp backs into the flower

and uses the claspers of the abdô-

men to attach itself to the base

of the lip. It then arches its body

and the apex of the abdômen

comes in contact with the visci-

dium and the stigma of the flower.

If the wasp is carrying pollinia

they are deposited on the stigma.

A certain externai similarity be-

tween the flower and the wasp

exist but the primary attractant is

the fragrance. The wasp has been

seen to eject seminal fluid and even

“preferred” the flowers to the real

females. Watson (1961) included

fine photographs in a short paper

on the same group.

Diuris is a genus of Australian

terrestrial orchids of which Cole-

man (1932) reported the pollina-

tion of D. pedunculata and D. sul-

phurea. The bees force the lip of

the flower back from the column

in order to reach the néctar ring

at their common base. The néctar

apparently acts as an intoxicant,

for some bees were observed with

their mandibles between the lip

and column for hours. Coleman

(1933) believed that both species

of Diuris are pollinated by male

bees stimulated in a similar

manner to Lissopimpla in the pol-

lination of Cryptostylis. The bees,

Halictus languinosus and Paracol-

letes sp., when landing on the lip

of the flower, curve the abdômen

so that its apex is extended slight-

ly over the apex of the lip (Ray-

MENT, 1932).

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Volume 5 (Zoologia)

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The flowers of Trichoceros and

its two closely allied genera, Teli-

pogon and Stellilabium, simulate

flies to a remarkable degree. Tri-

choceros is a genus of the Andes

of western South American. In

1962 I observed male Tachinid flies

of Paragymnomma sp. pollinating

flowers of Trichoceros antennifera

(Dodson, 1962 a). The column and

base of the lip are covered with

spiny, branched hairs as is the

abdómen of the flies. The lateral

lobes are narrow, barred with

yellow and redbrown and extend

laterally to similate the extended

wings of a sitting fly. The blade

of the lip has no particular similar-

ity to the head and thorax of the

fly but this probably is not neces-

sary to complete the illusion. The

stigma of the flower is located at

the apex of the “abdómen” of the

flower and reflects sunlight as does

the genital orifice of the female fly

when she opens and closes it to

attract the male. The viscidium,

extended over the stigma on

the slender rostellum, projects up

through the bristles and becomes

attached to the basal portion of

the fly. The male flies, deceived

by the similarity to the female fly

and stimulated by the signal from

the “genital-orifice-like” stigma,

land on the flower for an instant

(Fig. 30) and then pass on the

other flowers in the area. The pol-

linarium is deposited in a succed-

ing flower.

Fig. 30 — A male fly, Paragymnomma sp., attempting to copulate with the flower

of Trichoceros antennifera.

40

Atas do Simpósio sôbre a Biota Amazônica

Though we have no observations

of pollination of Telipogcn or Stel-

lilabium there seems no reason to

doubt that they are pollinated in

a similar manner.

Trigonidium occurs throughout

the lowland areas of much of trop-

ical America. Kerr & Lopez

(1963) reported pollination of Tri-

gonidium obtusum by male bees of

Trigona doryana in Brasil (Fig. 31).

The bees attempt to copulate with

the ápices of the petals, which have

conspicuous pad-like glands. A si-

milar gland occurs on the lip and

it is assumed that the bees enter

the flowers and attempt to copulate

with the lip as well.

Closely allied to Trigonidium are

the genera Mormolyca and Cyrti-

dium and their flowers are much

more insect-like. The column

. V

\

v

Fig. 31 — Male bees of Trigona doryana on the flower of Trigonidium obtusum.

(Sketch adapted from drawing forwarded by Dr. Sakagami) .

arches over the lip which usually

has a glandular-spot similar to

that of Trigonidium. It is assumed

that members of these two genera

are also adapted to pseudocopula-

tion as a means of cross-polination.

Pseudocopulation as a phenome-

non seems to be widely spread in

the orchids and may be preceeded

by sexual attraction through odors

which simulate those produced by

female insects ready for mating.

Vogel (1963) believed that the

attraction in Catasetum , Stanho-

pea and its allies is a part of a cycle

of pseudocopulation and that the

bees scratch on the lip out of frus-

tration. Sexual behavior may be

involved in the attraction of male

bees of the Euglossini through

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Volume 5 ( Zoologia )

41

strong fragrances but I feel that

there is much more to the story.

The odors produced may attract

the bees initially but the intoxica-

tion is fundamental to the opera-

tion of the complex pollination me-

chanisms in these orehids. The bees

must be drugged and must remain

for prolonged periods at the flow-

ers in order to pollinate them suc-

cessfully.

Pseudo-antagonism — This phen-

omenon was reported in Onci-

dium by Dodson and Frymire . . .

(1961 b) and corroborated by

observations of Dressler (personal

communication) . Male bees of the

genus Centris were observed at-

tacking the flowers of Oncidium

hyphaematicum and O. planilabre

in the Coastal zone of Ecuador. In

Centris the male bees are territo-

rial and drive away any other in-

sects that passes through. The

bees sit on sticks or leaves from

which they can survey their ter-

ritory. The racemes of the Onci-

dium flowers are long and arched

and move in the slightest breeze.

When the flowers move the bees

attack them if they are in his ter-

ritory. If the breeze continues the

bee attacks flower after flower.

Dressler reports that the territo-

riality is so fixed that the bees

return to their territories even

after being captured and released.

The bees do not land on the flow-

ers but simply strike them. The

viscidium attaches to the frons of

the bee’s head between the com-

pound eyes (Fig. 32) , and the stipe

rapidly depresses holding the pol-

linia extended directly in front of

the bee. Only certain species of On-

cidium have adapted to this be-

havior on the part of male Centris.

Other species are pollinated by fe-

male Centris which attempt to col-

lect néctar from the flowers. The

callus apparently functions to po-

sition the female bee properly

when she lands.

Pseudoprey — This is an unprov-

en category. Fordham (1946) re-

ported a scoliid wasp, Campsomeris

tasmaniensis , making stinging mo-

vements on the labellum of Calo-

chilus campestris in Australia.

Unfortunately, the sex of the pol-

linator was not determined but if

it had a sting it was a female. Fe-

males of this group lay their eggs

on the larvae of other insects and

when the eggs hatch the wasp

larvae consumes the body of the

host.

EVOLUTION IN ORCHID

FLOWERS

The orchid flower is highly spe-

cialized for insect pollination. The

very characteristics which act to

circumscribe the orchid family

tend to be adaptations to special-

ized pollinator-flower relationships.

The most primitive of living or-

chids are well along the evolution-

42

Atas do Simpósio sôbre a Biota Amazônica

Fig. 32 — Male Centris geminata attacking the flower of Oncidium planüabre

attempting to drive the insect-mimicing flower away. Note the pollinarium

attached to the head of the bee.

ary path to pollinator specificity.

The typical orchid flower has one

petal developed into a lip for a

landing platform. The styles and

anthers are joined into a single

structure, the column, which is

designed to place the pollen on the

pollinator at a specific place. The

pollen is gathered into masses cal-

led pollinia and is arranged so that

pollinators are unable to collect it

for food. Tremendous numbers of

pollen grains in each pollinium fer-

tilize similarly great numbers of

ovules in the ovary when pollin-

ation does occur. Literally millions

of tiny, wind-distributed seeds can

be produced in each ovary. The

seeds have no endosperm and con-

sist of a few cells surrounded by a

papery cover. They are so small

and light that they can be carried

great distances by the wind, but

in order to germinate they normal-

ly must be invaded by a fungus.

THE ANCESTRAL ORCHID FLOWER AND

ITS POLLINATORS

The ancestral orchids were prob-

ably pollinated by bees or wasps.

It seems likely that they simply

produced néctar inside néctar

tubes developed from the sepals

and that the bees did not collect

pollen. The flower was probably

cupped and zygomorphic and the

bees landed on a petal which was

enlarged to form a landing plat-

Volume 5 (Zoologia)

43

form. No present day flower could

be considered as similar, though

the members of the subfamily

Apostasioideae are only slightly

advanced. When the pollinator

attempted to reach the néctar the

granular pollen was brushed onto

its body from one of the three or

Fig. 33 — Hypothetical ancestoral orchid flower.

more anthers. On visiting a suc-

ceeding flower the pollen was de-

posited on the stigma and a new

load of pollen was received. Ag-

gregation and reduction of the

stamens and pistil to form a

column and its refinements have

made the system more efficient.

Additional refinements in trans-

port mechanisms of pollen masses

44

Atas do Simpósio sôbre a Biota Amazônica

and in attraction of more specific

pollinators provided for develop-

ment of abundant species but the

basic system remains unchanged.

In Figure 33 a schematic diagram

depicts my idea of an ancestral or-

chid flower (see also Dressler &

Dodson, 1960 and Garay, 1960).

TRENDS IN FLORAL ADVAN CEMENT AND

SPECIALIZATION FOR POLLINATION

From the ancestral type of or-

chid flower two lines of develop-

ment were successful. They were

primarily expressed in consolida-

tion of the column into two or

three anthers on one hand or to

a single anther on the other. The

Apostasioideae still retain two or

three anthers and form a very

small group of species in two ge-

nera. The Cypripedioideae have a

column with an anther on each

side. The third anther forms a

broad, sterile, plate-like staminode

at the apex of the column while the

stigrna is a broad plate on the un-

derside of the column. The group

is typified by having one petal

formed into a cup-like trap into

which the pollinators fali and must

pass under the stigma and an

anther in order to escape. About

100 species form four genera.

The other major line, with a

single anther, forms three natural

sub-families; the Neottioideae, the

Orchidoideae and Epidendroideae

(Dressler & Dodson, 1960 and

Dressler, 1962) . These subfamilies

contain thousands of species in

nearly 600 genera. The Neottioi-

deae are terrestrial orchids usually

found in the temperate zones. They

have soft, mealy pollinia and the

stems of the plants do not form

corms or other thickenings. The

Orchidoideae are also temperate

zone terrestrial orchids with soft

pollinia but here the pollinia have

caudicles arising from their base.

The caudicles are formed from

pollen and strands of viscin and

aid in attachment to the pollin-

ator. These two subfamilies are not

large (perhaps with 2,000 species)

whereas the Epidendroideae con-

sists of from 15,000 to 20,000 spe-

cies in more than 500 genera. The

pollinia are usually hard and waxy

and if they have caudicles they

arise from the apex. Most of the

species are found in tropical re-

gions and the majority live as epi-

phytes.

It is in the Epidendroideae

that advancements in adaptations

based on pollinator phenomena

have developed to their most ad-

vanced state. Here placement of

the pollinia on the pollinator be-

comes important and mechanical

devices for attaching the pollinia

become complex. The simple Sys-

tem found in the lower orchids, in

which the pollinator contacts the

sticky stigma with its dorsal sur-

face and the pollinia are attached

Volume 5 (Zoologia)

45

as the pollinator backs out of the

flower (Fig. 34 a) , is still found in

the less advanced forms in the Epi-

dendroideae. However, the pollinia

often have caudicles which become

attached to the pollinator. In some

large and relatively advanced

groups such as the Dendrobium,

Bulbophyllum and Pleurothallis

allies no caudicles are found and

the pollinia simply become attach-

ed to the sticky surface of the

pollinator. In the Epidendrum al-

liances caudicles are well developed

(Fig. 34 b).

A further advance is the visci-

dium — a piece of the rostellum

separating the anther from the

stigma — which comes away as

the pollinator pushes against it

(Fig. 34 d) . The pollinia are attach-

ed to the viscidium by viscin

threads and the side of the viscidi-

um toward the pollinator is sticky.

This is a fundamental advance

and is found in many genera such

as the Maxillaria and Cymbidium

allies. The addition of a piece of

sterile tissue between the viscidium

and pollinia, called the stipe, is

an even greater advance and is

found in the allies of Oncidium

and Vanda and in most of the

allies of Maxillaria though not in

Fig. 34 — The column and arrangement of pollinia and accessory organs. A,

Cephalanthera. B, Cattleya. C, Oncidium. D, Cymbidium.

46

Atas do Simpósio sôbre a Biota Amazônica

Fig. 35 — Drawing of the pollination mechanism of a typical bee-pollinated

orchid flower. A, Cross-section of flower. B, Bee with pollinarium from another

flower on its thorax entering the flower. C, Bee Beginning to withdraw from the

flower. D, Pollinarium on the back of the bee is deposited on the stigma as the

rostellum engages the back of the thorax of the bee. E, Bee forcing past the

rostellum and against the anther. F, Rostellum passing over the thorax of the

bee depositing sticky fluld. The pollinia in the anther are attached to the bee by

the sticky fluid from the stigma. G, The bee prepared to fly to another flower.

Volume 5 (Zoologia)

Maxillaria itself (Fig. 34 c) . The

stipe allows for greater precision

in insertion of the pollen masses

into the stigma. Also, through dif-

ferential drying of the outer and

inner surface, the relative position

of the pollinia in reference to the

anther and the stigma can be

changed (See Darwin, 1862 for an

extensive discussion of this pheno-

menon) . The total unit of pollinia,

viscidium, caudicles, or stipe is

called the “pollinarium”.

The advancement tends to be

from naked, soft masses of pollen,

47

to soft masses with caudicles, to

waxy masses with or without cau-

dicles, to pollinia attached to a vis-

cidium and finally to a viscidium,

stipe and pollinarium.

The placement of the pollinia on

the body of the pollinator also

shows trends in specialization. The

most common and basic system in

the orchids is that shown in

Fig. 35 in which the pollinia

are placed on the dorsal surface

of the thorax of the pollinator. As

the attachment devices become

more complex, placement at other

Fig. 36 — Morphological reorganizations in bee pollinated orchids for placement

of pollinaria on different parts of the bee. A, Typical bee-pollinated orchid flower

where pollinaria are deposited on the thorax of the bee (Cattleya). B, Pollinarium

deposited on the front of the head (Oncidium). B1, Pollinarium attached to head

of Bee. B\ Stipe of pollinarium depressing. B3, Pollinia deposited in stigma while

a new pollinarium is attached to the head. C, Pollinarium attached to the back

of the head of the bee.

cm

48

Atas do Simpósio sôbre a Biota Amazônica

spots on the body of the pollinator

become possible. When the pollinia

are placed on the back of the head

the column is similar but shorter.

When pollinia are placed on the

front of the head of the bee an

abrupt change occurs and the co-

lumn is erect with viscidium cr

caudicles twisted around and fac-

ing outward so as to contact the

head of an insect forcing inward,

rather than facing toward the in-

side of the flower and being attach-

ed to an insect backing out (Fi-

gure 36) .

Naturally in butterfly-, moth-

and bird-flowers the pollinia are

placed on the proboscis or beak.

Here the same system of develop-

ment of pollinaria complexities

hold but these pollinators are not

basic and adaptations are required

to make basically bee-flowers func-

tion efficiently when pollinated by

an animal with a proboscis or bill.

These adaptations are discussed

below. Special adaptations such as

hooked viscidia and elongated

stipes make it possible to place pol-

linia on the legs of the pollinator

and occur in such highly specializ-

ed genera as Haemeria, Trichoce-

ros, Sievekingia and Ornithoce-

phaus. Differential tension on the

stipe makes possible the pollen-

-throwing system found in the sub-

tribe Catasetinae.

MORPHOLOGICAL COMPENSATIONS IN

THE FLOWER TO DIFFERENT KINDS

OF POLLINATORS

The morphological compensa-

tions in the basically bee-pollin-

ation oriented orchid flower to dif-

ferent kinds of pollinators directly

reflects the visiting and feeding

habits of those animais. For the

case of moths and butterflies the

lip of the flower is usually joined

to the column to form a long

néctar tube (Fig. 37). In addition

the rostellum is cleft so as to guide

the proboscis properly into place so

that the pollinia can be attached

easily at a position which will

make deposition on the stigma

easy on the next flower visited. For

butterfly-flowers the lip or the

whole cluster of flowers are ex-

tended to form a landing platform.

In moth-flowers the lip is usually

reduced or curled back out of the

way.

Bird-flowers may be of two types

depending on their immediate an-

cestors. If bird-flowers are derived

from butterfly-pollinated ancestors

they will closely resemble butter-

fly-flowers. However, if they have

immediate bee- or fly-pollinated

ancestors strong compensations

are needed to force the beak

against the column for pollinia de-

position. In most cases this is ac-

complished by calli on the lip of

the flower which simply force the

beak upwards as it is introduced

Volume 5 (Zoologia)

49

Pig. 37 — Orchid flowers adapted to pollination by other agents. A, Butterfly-

-pollinated Epidendrum. B, Moth-pollinated Epidenãrum. C, Bird-pollinated

Epidendrum derived from either Lepidoptera-pollinated ancestor. D, Bee-

-pollinated Rodriguezia leeana. E, Bird-pollinated Rodriguezia secunda.

until it contacts the pollinia (Fi-

gure 38) .

Fly-flowers are often very simi-

lar to bee-pollinated flowers. How-

ever, in most cases additional

guidance mechanisms are neces-

sary to entice or force erratic flies

to act as efficient pollinators.

Often the sepals and petals are dev-

eloped into long “tails” or may be

joined to form a radial, saucer-

-like flower or cluster of flowers.

Tassels and flexible or articulated

bracts or other devices which move

in the slightest breeze act to at-

tract the attention of the fly. Color

schemes also contribute to the

attraction. Finally, once the fly has

landed and proceeds to investigate

the flower of many fly-pollinated

orchids, the lip, which is either ba-

lanced on a pedestal or held under

tension, throws the insect against

the column and pins it there.

Usually after a considerable strug-

gle — during which the pollinia

4 — 37 145

50 Atas do Simpósio sobre a Biota Amazônica

are cemented in place or deposited

on the stigma — the fly escapes

to go on to another flower. These

mechanisms are common in Bulbo-

phyllum and Plocoglottis. Some or-

chids such as Pterostylis, produce

traps which imprison the flies

until they have had time to pick

up the pollinia. Cryptophoranthus

produce a lantern-like flower with

slits in the sides of the joined se-

pals for the entrance of the pol-

linator similar to that found in Ce-

ropegia of the Asclepiadacea. Most

members of Pleurothallis and Mas-

devallia have rather simple mecha-

Fig. 38 — Morphological characters of bird-pollinated orchids. A, An orchid

derived from bee-pollinated ancestors, Rodriguezia seconda, with the lip close to

the column to force the beak upward. B, Epidendrum derived from butterfly-

-pollinated ancestors with callus of the lip forcing the beak of the bird upward

against the column.

cm

Volume 5 (Zoologia)

51

nical systems but one species of

Masdevalia, M. muscosa has a sen-

sitive lip which probably carries

the pollinator as in Bulbophyllum.

These adaptations from a rela-

tively simple bee-flower to mechan-

isms facilitating pollination by

other insects and birds are cha-

racteristic for particular groups

and have played an important role

in orchid classification on the gen-

eric levei . Almost universally

however, taxonomists have not un-

derstood either the mechanisms or

their importance and have simply

grouped plants which have similar

characteristics into genera regard-

less of the origin or importances

of those salient characters. In

small groups few major mistakes

have been made, but in such large

groups as the Pleurothallis, Bulbo-

phyllum, Epidendrum and Onci-

dium complexes, chãos is the re-

sult. Until taxonomists understand

why these plants are similar or dis-

tinct, chãos will continue to reign.

Additional variation and problems

have resulted from adaptations to

different agents at later times, so

that moth or fly-pollinated or-

chids, for instance, may be primi-

tive or advanced, may characterize

whole subtribes or only isolated

species in large genera adapted

otherwise, or may form part of a

varied spectrum of pollinators in a

constellation of small, closely al-

lied genera.

IMPORTANCE OF POLLINATOR SPECIFI-

CITY IN SPECIATION IN THE ORCHIDS

Grant (1963, ch. 13) has writ-

ten a lucid account of isolation

mechanisms in which (following

Dobzhansky, 1951) he has pointed

out: “Two or more biotically sym-

patric populations — must be re-

productively isolated in order to

maintain their separate genetic

constitutions. Reproductive isola-

tion is a characteristic of species”.

Orchids follow the general pattern

of other plant groups in respect to

isolation of species with the same

three isolating factors, i.e., 1. Spa-

tial, 2. Environmental, and 3. Re-

productive, playing their normal

roles. However, the orchids, as a

result of their intrinsic nature,

have been more heavily oriented

toward that of reproductive isola-

tion. Grant and others divide re-

productive isolation into two major

divisions: externai and internai

factors. ^-oth factors play impor-

tant roles in the Orchidaceae but

by far the more important is that

of externai reproductive isolation

which Grant further divides into

mechanical, ethological (behavior

characteristics), and seasonal iso-

lation.

Species-separation in the orchids

is primarily based on a combina-

tion of mechanical and ethological

isolation. The flower may attract

several different pollinators, only

one of which is able effectively to

52

Atas do Simpósio sôbre a Biota Amazônica

pollinate the flower due to mechan-

ical exclusion of the other pollin-

ators (Dobzhansky, 1937; Steb-

bins, 1950; and Dodson, 1962 a),

e.g., Stanhopea florida and S. con-

nata, which are sympatric in east-

ern Ecuador. Both species attract

Euglossa nigropilosa but in the

case of S. connata this bee is not

large enough to effect pollination

whereas the larger Eulaema spe-

ciosa is also attracted and does

pollinate the species. No hybrids

occur in nature but are easily

made artificially. Near Turrialba

in Costa Rica I found six species

of Stanhopea to be sympatric,

5. costaricensis, S. gibbosa, S.

pulla, S. ecornuta, S. wardii and

S. warszewicziana. Stanhopea cos-

taricensis and S. gibbosa hybridize

frequently (Fig. 39) because their

two pollinators are very closely

allied and occasionally make

errors. The result is a vast hybrid

swarm. Stanhopea costaricensis

Fig. 39 — A hybrid between Stanhopea costaricensis and Stanhopea gibbosa

being pollinated by Eulaema meriana in Costa Rica.

(Fig. 40) and S. ecornuta (Fig. 41)

very rarely hybridize. Stanhopea

costaricensis is pollinated by Eu-

laema seábrai while S. ecornuta is

pollinated by Euplusia schmidtia-

na. I have, however, observed Eu-

laema seabrai visiting S. ecornuta

and if such a bee were carrying pol-

linia of S. costaricensis hybrids

could occur. The pollinia of S. ecor-

nuta are too large to be inserted

in the stigmatic pocket of S. cos-

Volume 5 (Zoologia)

Fig. 40 — Stanhopea costaricensis being pollinated by Eulaema meriana.

Fig. 41 — Stanhopea ecornuta.

cm

54

Atas do Simpósio sôbre a Biota Amazônica

taricensis and therefore the cross

could only occur in one direction,

i.e. S. ecornuta as the pod parent.

A few plants have been collected

which were unquestionably hybrids

between these two species. They

apparently are not visited, how-

ever for they remain very rare and

no evidence of back-crossing with

either parent has been found. The

hybrids appear to be reproductive-

ly sterile and in terms of evolu-

tion are merely an artifact. If, how-

ever, an effective pollinator ap-

peared they could lead to a new

evolutionary line in Stanhopea. A

hybrid between S. ecornuta and

S. graveolens was found in Guate-

mala and was named 5. Lewisae

by Ames and Correll. The other

species at Turrialba are maintain-

ed as separate populations by pol-

linator specificity.

On the other land, the flower

may attract only a single kind of

pollinator by acting upon the be-

havioral characteristics of that

animal (Dodson & Frymire, 1961b

and Dodson, 1962a), for instance,

two species of hummingbird are

attracted to two differently color-

ed sympatric species of Elleanthus

in Ecuador. Different species of

Euglossa are attracted to different

forms of Gongora maculata (mak-

ing them biological species at

least) through the production of

different odors (Dodson, 1965 and

Dressler, personal communica-

tion) .

On the basis of such information

I postulated (Dodson, 1962 a) a

possible scheme of speciation in

Stanhopea which I (with tongue-

-in-cheek) called “leap-frog specia-

tion”, whereby species could devel-

op in the genus by small, spatially-

-isolated populations simply adapt-

ing to differently sized pollinators.

The populations of newly developed

species could later become sympa-

tric but would be reproductively

isolated. A prime example is that

of Stanhopea florida and S. nigri -

pes, which have large populations

in Peru that are only sympatric in

a relatively small area and probab-

ly recently have become so. The

habitat, habit of the plants, and

the flowers of the species (Fig 42) ,

are nearly identical except that

S. florida is pollinated by a small

Euglossa while S. nigripes is pol-

linated by a much larger Eulaema.

Though the flowers are nearly

identical in size and form, they

differ in color, size of the exit

space, and odor produced. No na-

tural hybrids have been found. The

two species are now clearly distinct

but are probably recently derived.

Conceivably further speciation can

easily occur in these orchids

through other isolated populations

being adapted to a different species

of Euglossa or Eulaema.

Volume 5 (Zoologia)

55

Fig. 42 — Illustration of Stanhopea florida on ieft and Stanhopea nigripes on

right. The two flowers are very similar but though sympatric in Peru do not

hybridize naturally due to adaptation to different-sized bees.

Similar phenomena are evident

in other highly pollinator-specia-

lized orchids and are in the pro-

cess of being documented.

Grant (1949) made an interest-

ing survey of plants belonging to

various genera and families of

flowering plants. He found that in

those groups pollinated by specia-

lized, flower-visiting animais, 37 to

54 percent of the taxonomic char-

acters used to separate related

species of non-promiscuous Angio-

sperms were involved with the co-

rolla, stamens and pistil. Converse-

ly, floral characteristics only

comprised 15% of the characters

used to distinguish related species

of plants pollinated by miscella-

neous insects. I estimate that in or-

chids approximately 90% of the

characters used to differentiate

species deal with floral characters.

In orchids with extremely intricate

floral mechanisms and high speci-

fity of pollinator attraction nearly

100% of differentiating characters

between species are within the

56

Atas do Simpósio sobre a Biota Amazônica

flower and more than 60% are in

the lip and column alone, whereas

in more simple orchids visited by

various kinds of bees, 77% of the

characters deal with the flower.

These figures clearly demons-

trate the importance of pollination

phenomena in a highly advaneed

and pollinator-oriented group such

as the orchids if only for the use

of taxonomists working with the

species. Obviously their importance

in speciation for the orchids is fun-

damental.

LIST OF ORCHIDS AND THEIR KNOWN POLLINATORS

Volume 5 (Zoologia)

57

Orchid

Bulbophyll,um

B. macranthum

B. striatellum

Caladenia

C. alba

C. barbossae

C. dejormis

C. dialata var.

rhomboidijormis

C. dirnorpha

Caleana

C. alba

Calochilus

C. campe&tris

Calopogon

C. barbatus

C. pulchellus

Calypso

C. borealis

Catasetum

C. barbatum

C. bicolor

C. cemuum

C. costatum

C. dilectum

C. discolor

C. eburneum

(as C. fragrans)

C. jimbriatum

C. hookeri

C. integerrimum

C. luridum

C. macrocarpum

58

Atas do Simpósio sôbre a Biota Amazônica

Volume 5 (Zoologia)

59

Cryptostylis

‘Nonpollinating visitors.

60

Atas do Simpósio sobre a Biota Amazônica

Volume 5 (Zoologia) 61

62

Atas do Simpósio sôbre a Biota Amazônica

•Nonpollinatmg visitors.

64

Atas do Simpósio sobre a Biota Amazônica

‘Nonpollinating visitors.

SciELO

10 11 12 13

Volume 5 (Zoologia)

65

5 — 37 145

66

Atas do Simpósio sôbre a Biota Amazônica

**Many other pollinators of members of this genus have been reported in European litera-

ture. We mainly cite the basic observations of Darwin, Muller and Robertson and therefore

adhere to their nomenclature here.

68

Atas do Simpósio sôbre a Biota Amazônica

•Xonpollinating visitors.

70

Atas do Simpósio sôbre a Biota Amazônica

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Ridley, H. N., 1890a, The genus Brom-

headia. Linn., J., 28: 331-339.

Ridley, H. N„ 1890b, On the method of

fertilization in Bulbophyllum ma-

cranthum and allied orchids. Ann.

Bot., 4: 327-336.

Ridley, H. N., 1896, Orchideae and

Apostasiaceae of the Malay Fen-

ninsula. J. Linn. Soc. Bot., 32: 213-

-416.

Ridley, H. N., 1905, On the fertilization

of Grammatophyllum. Roy. Asiatic

Soc. Str. Br., 44: 228-229.

Robertson, C., 1887, Fertilization of Ca-

lopogon parviflorus. Bot. Gaz., 12:

288-291.

Rogers, R. S., 1931, Pollination of Ca-

ladenia deformis. Trans. Roy. Soc.

S. Australia, 55: 143-146.

Rolfe, r. a., 1910, The bee-orchis.

Orch. Rei >., 10: 261.

Sargent, O. H„ 1909, Notes on the life-

-history of Pterostylis. Ann. Bot.,

23: 265-274.

Sargent, O. H., 1934, Pollination in Pte-

rostylis. Vict. Nat., 51: 82.

Slade, H., 1962, Some attractive Den-

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Orch. Soc. Buli., 31: 993-996.

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lution in plants. Columbia N.Y.

Thomson, G. M„ 1927, The pollination

of New Zealand flowers by birds

and insects. Trans. Proc. New Zea-

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Atas do Simpósio sôbre a Biota Amazônica

Vol. 5 (Zoologia): 73-83 — 1967

PREDATOR-PREY INTERACTION S AND THE EVOLUTION

OF ASPECT DIVERSITY

A. STANLEY RAND

Smithsonian Tropical Research Institute, Balboa, Canal Zone

The diversification of life is one

of the major trends in evolution

and one that reaches a maximum

in the tropics. Even within the tro-

pics some areas are richer than

others and every biologist who has

visited the Amazônia rain forests

has remarked the wealth of life

here in both number of species and

variety of different types or as-

pects. This conference on the biota

of the Amazon seems an appro-

priate place to discus an idea about

the evolution of diversity.

Richness of tropical life has

many facets, some of them of prac-

tical importance in tropical agri-

culture and medicine, and all of

species co-existing in an area, is

more or less theoretical interest.

Species diversity, the number of

the problem currently receiving

most attention and the associated

hypotheses, theories and concepts

have been well reviewed recently

by PlANTCA (1966).

However, there is another strik-

ing attribute of the life in tropic-

al rain forests, the existence ol

many varied and extreme types,

bazaar forms and improbable co-

lors (Kettlewell, 1959). In this

paper I want to discuss the evolu-

tion of these differences in appear-

ance that for convenience. I will

call aspect diversity.

I was first impressed with the

problem of aspect diversity while

watching moths around the lights

at the Estação Biológica de Bora-

céia of the Departamento de Zoo-

logia, São Paulo. Not only were

there a fantastic number of species

but they were strikingly different

from one another and these differ-

ences were accentuated by the va-

riety of positions taken by the rest-

ing animais. I have continued to

watch insects on Barro Colorado

Island where I am now working

and am convinced that the differ-

74

Atas do Simpósio sobre a Biota Amazônica

ences in appearance, i.e. the differ-

ences themselves, must be adapt-

ive.

Predators are an important se-

lective force on the appearance of

their prey as Cott (1940), Kettle-

well (1961), and Cain & Shep-

pard (1950) have shown conclusive-

ly. I think predators have been

an important factor in the evolu-

tion of aspect diversity.

I would like to suggest the fol-

lowing two part hypothesis: A. Be-

cause learned characteristics play

an important role in prey recogni-

tion by visually oriented verte-

brates there is a selection pressure

on their prey species to differ from

one another in appearance. B. The

effects of this selection pressure

will be greatest where predator-

-prey relationships are most com-

plex.

Uexküll (1957) in this provoc-

ative discussion of Umwelts pointed

out that people form search im-

ages, i.e. one finds what one is look-

ing for, what matches one’s

expectation. Sometimes one can

look directly at an object being

searched for and not “see” it be-

cause it is not the expected shape

or color.

The formation of a search image

is a familiar experience to every

naturalist. The first specimen of

any species is always the most dif-

ficult to find and after the first

few have been located an even ap-

parently uncommon species may

be seen everywhere.

L. Tinbergen (1960) first em-

phasized the role of learning or for-

mation of a search image in prey

recognition by wild birds. They im-

prove with practice their ability to

find cryptic insects and each dif-

ferent kind must be learned sepa-

rately.

Ruiter (1952) showedthat naive

Jays ( Garrulus glandarius) and

wild caught adult Chaffinches

( Fringilla coelebs) presented with

a mixture of sticks and stick ca-

terpillars took a long time to find

the first stick caterpillars, but

quickly found the rest. Once the

first Caterpillar had been found

half the Jays pecked at both the

sticks and caterpillars that they

had been ignoring completely. The

other Jays found the caterpillars

but either did not pick up sticks

at all or only did so after they had

found and eaten all the caterpil-

lars. Those that did pick up sticks

picked up only those that re-

sembled the caterpillars and not

the other types. N. Tinbergen. . . .

(1957: 52) in discussing these ex-

periments further records that

“When the caterpillars outnumber-

ed twigs, the birds took the disap-

pointments in stride and went on

hunting for caterpillars. But if

they picked up more twigs than

caterpillars, they became discour-

aged and gave up searching.”

Volume 5 (Zoologia)

75

Insect eating mammals also im-

prove their prey finding with prac-

tice. Recent experiments by M. Ro-

binson on Barro Colorado Island

have shown that naive captive

marmosets at first ignored stick-

-like phasmids. But once a marmo-

set had found one it very quickly

found others, making discrimina-

tions between stick and insect that

it had not previously made. When

the marmosets were presented with

only sticks for a series of trials they

again ignored the stick insects.

L. Tinbergen (1960) presented

evidence that the formation of a

search image may be important in

nature. In a pinewood in Holland

he studied the variations in the

number of each species of insect

that Great Tits were bringing to

their nestlings and the relation-

ships of these to variations in the

abundance of the prey species in

the areas being searched by the

birds. The proportion of the indi-

viduais present in the area that

were being taken by the Tits was

very low when a species was rare

and rose sharply to a much higher

proportion when the species be-

came moderately common. He sug-

gested that the chance encounters

between Tit and insect are too in-

frequent at low prey population

densities to allow the Tits to form

effective search images. When the

prey becomes more common the

chance encounters increase and

the Tit learns to recognize the in-

sect and find individuais that it

would earlier have missed.

Tits never concentrated on only

a single prey item even when it

was very abundant; instead they

apparently use several search im-

ages either simultaneously or in

quick succession.

The number of search images

that can be retained and the effect

of learning a new search image on

those already learned is an im-

portant aspect of predatory beha-

vior yet unexplored.

Mook et alii (1960), working

on the same experimental situa-

tion, showed that there was a lag,

sometimes of several days, between

the time when a cryptic prey spe-

cies increased in abundance and

the time when Tits began to feed

on it heavily. This lag varied from

pair to pair but seldom between

the two members of a pair. This is

further evidence that it takes sev-

eral chance encounters before a

Tit learns to hunt a new prey spe-

cies efficiently.

Though the experimental work

so far has dealt with only birds

and mammals, something like the

formation of a search image occurs

in other vertebrates. The trout in

a stretch of stream sometimes will

take one type of lure and some-

times another. Usually the lure

taken imitates something on which

they are already feeding. It would

76

Atas do Simpósio sobre a Biota Amazônica

be interesting to know if this in-

volves the formation of search im-

ages like that in birds and if so how

general this phenomenon is among

predators.

The role of learning in the re-

cognition of warningly colored in-

sects and the evolution of mimicry

has been investigated much more

thoroughly by the Browers (e.g.

1960 and 1965), Sexton (1960),

and others, than has the role of

learning in finding cryptic prey.

It is tempting but unwise to as-

sume that the patterns of learning

which are exhibited in the mimicry

studies will hold in the recognition

of cryptic prey. Certainly the stu-

dies on imprinting show that the

same animal may learn different

kinds of things in very different

ways and that speeds of learning

and forgetting vary with age and

situation.

One very important result of the

work of L. Tinbergen (1960) and

Mook et alii (1960) that should be

investigated further is the exist-

ence of a threshold at which chance

encounters are frequent enough to

cause the formation of a search

image. This implies that changes

in prey density across this thresh-

old seem to have much more effect

on degree of predation than com-

parable changes in density above

and below the threshold.

A threshold effect may occur in

certain situation or in certain pre-

dators while in others the ability to

find a cryptic insect may increase

more uniformly as the frequency

of contacts rises.

In any case a cryptic insect seems

least likely to be found by a

vertebrate predator that has never

caught one of its sort before and

most likely to be found by a preda-

tor that has just caught several.

The unique species is at a selec-

tive advantage. If two species are

sufficiently similar in appearance,

experience with either is useful to

a predator in finding the other.

This is therefore clearly a disad-

vantage to both prey species for it

increases the opportunities of a

predator to form a search image

and this will increase the predator’s

efficiency in hunting for both spe-

cies. Since the differences between

the two species will be of selective

advantage to both they will tend

to become more and more different

from each other and from all other

species preyed on by the same pre-

dators. Selection will be for aspect

diversity. It will primarily involve

edible species that depend on not

being seen or recognized as food for

their major line of defense against

predators. Distasteful species will

tend to become more conspicuous

and more similar to one another

making the predators’ recognition

of them easier. Thus Müllerian

mimicry is in one way the opposite

of aspect diversity.

Volume 5 (Zoologia)

77

Obviously a species gains an ad-

vantage from being unique in ap-

pearance only when it is one of sev-

eral prey of a polyphagus preda-

tor. Any species that is heavily

preyed upon by a monophagus pre-

dator that hunts by sight and a

search image may lose rather than

gain by being unique.

From our hypothesis one would

expect animais in the same habitat

to differ in appearance even when

they are using the same kinds

of camouflage. Herrebout et alii

(1963) report that the caterpillars

that live together among pine

needles in Dutch woodlands show

exactly this. There are six species

with a green and white striped

p a 1 1 e r n that is demonstrably

cryptic but no two species have the

stripes arranged in the same way.

The authors interpret this as in-

complete convergence of distantly

related caterpillars. It surely is this

but I would suggest the very in-

completeness, i.e. the observed

differences between species, are

themselves adaptive.

Preliminary observations on in-

sects on Barro Colorado Island

suggest the same phenomenon.

Sphyngid moths (about 20 species)

and leaf-like Katydids (13 species)

were collected around the labora-

tory lights for several nights. In

both the Sphyngids and the Katy-

dids the same basic type is present

throughout but each species shows

striking visible differences from

the others.

I also collected samples of or-

thoptera living in the grassy clea-

ring around the laboratory. Here

several different stocks have evolv-

ed patterns that are basically

green and cryptic in the grass but

the eight species represented are

still very different from one ano-

ther. In general aspect the differ-

ences within families are as great

as those between families.

These observations fit the sug-

gested hypothesis. Though they do

not prove it they demonstrate the

reality of the problem. Testing the

hypothesis must be done experi-

mentally; observations of the sort

just reported serve only to indicate

that the experiments are worth

doing.

There is one alternate hypothe-

sis that must be considered: that

the diversity in aspect in tropical

species of cryptic animais is be-

cause each is matching a slightly

different background. The work on

caterpillars in Holland and my

observations on Barro Colorado Is-

land suggest that this is not true

but the possibility needs rigorous

testing.

L. Tinbergen’s studies suggest

that the effect of being different

from other prey species may be

most important at relatively low

population densities. Once a spe-

cies is common enough that

78

Atas do Simpósio sôbre a Biota Amazônica

chance encounters with it alone

are enough to lead to a search

image another strategy, variability

within the species, becomes im-

portant. Sheppard (1958) has dis-

cussed this point and the further

one that a predator feeding pre-

ferentially on the commonest

morph could act to maintain po-

lymorphism in a population. The

only case I know where this lat-

ter hypothesis has been seriously

considered is in the snail Cepia

nemoralis and here it seems theo-

retically possible but not an im-

portant mechanism (Cain & Shep-

pard, 1954; Ford, 1964) .

Brower (1958) discussed an-

other way prey species may react

to predators that form a search

image. He suggests that originally

polyphagus caterpillars with simil-

ar appearance might specialize in

different food plants and so reduce

the opportunities for a predator to

form an effective search image.

A further possible consequence

of the search image formation is

that it may act to promote species

diversity. Paine (1966: 65) advan-

ced the hypothesis that “Local

species diversity is directly related

to the efficiency with which pre-

dators prevent the monopolization

of the major environmental requis-

ites by one species.” Based on

studies of intertidal marine organ-

isms the suggestion may have

wider applications and be par-

ticularly important in situations

where visually hunting vertebrates

are major predators.

The discussion so far has been

of prey recognition by birds and

other vertebrates. Certainly many

predators, in the wide sense, are

insects in which learning plays a

much more limited role (Thorpe,

1956) . For example, mantids (Ril-

ling et alii, 1959) seem to recognize

only moving prey as food and to

be most attracted by objects with

projections that move indepen-

dently of the main moving mass.

They can learn to ignore an object

but don’t seem to learn to attack

one.

A related sort of prey recogni-

tion also occurs in vertebrates and

experiments with cryptic animais

has shown that movement is one

of the most important cues used.

Cott (1940) presents cogent ar-

guments that eyes are another ge-

neral cue. In these cases the pre-

dator responds to characteristics

that the prey all have in common.

In invertebrates what cues are

used is largely genetically deter-

mined. In vertebrates learning may

also play an important role.

Though this system of prey re-

cognition is very different from the

formation of a search image, its

effect on the aspect diversity of the

prey will be somewhat similar. Any

prey that can change its appear-

ance so that it no longer exhibits

Volume 5 (Zoologia)

79

the relevant cues will be ignored.

If many species change in the same

way the old cues will be abandon-

ed and others adopted. Thus se-

lection will favor the unique spe-

cies and aspect diversity will be

increased.

If the first section of the hypo-

thesis, that the formation of a

search image leads to aspect diver-

sification, is true, it seems evident

that the greatest effect of this will

occur where the predator-prey in-

teractions are most complex, i.e.

where the number of prey alterna-

tives for each predator species and

the number of predator species per

prey species are greatest.

The more prey species included

in the diet a predator the more di-

vergent and extreme at least some

must be if all are to be unique.

Cryptic species can be similar

in appearance only when all are

so rare that chance encounters are

not common enough with any of

them to allow formation of an ef-

fective search image. The more

species there are in any such group

the more likely that one will in-

crease to the point where a search

image covering all is formed and

strong selection for divergence

produced. The more kinds of pre-

dators there are in a system the

more kinds of search images can

be formed and consequently the

differences between prey will tend

to be either more extreme or more

numerous or more detailed.

Thus it seems that where the

predator-prey relations are most

complex, aspect diversity will be

greatest. Certainly tropical areas,

particularly rain forests, have

large numbers of prey and of pre-

dator species.

It can be argued that in tropical

regions predators are more special-

ized than they are in temperate

regions and therefore predator-

-prey relationships not more

complex even though more species

are present. The amount of special-

ization in tropical animais is stili

being studied (Klopfer & MacAr-

thur, 1960) but it is clear that

tropical birds, however specialized

in their feeding niches, stili take

a variety of food. Skutch (1954);

1960) lists the food of 50 species

of Central American birds that

eat insects. Of these he records

that 29 also eat vegetable material

and he saw an additional 12 eat

animais other than insects. No bird

restricted its diet to a single or a

few prey species. As a completely

different example: Inger & Marx

(1961) analyzed the food of 20

species of amphibians from the

Congo. All 20 had eaten primarily

insects but all had eaten other

animais as well. Within the insects

each species of amphibian had

eaten representatives of at least

six (usually 9 to 11) orders. How-

80

Atas do Simpósio sobre a Biota Amazônica

ever specialized tropical predators

may be the vertebrates seem poly-

phagus enough to exert a selection

for aspect diversity.

In this paper I have discussed

certain aspects of diversity in

terms of the hypothesis that: Be-

cause learned characteristics play

an important role in prey recogni-

tion by visually oriented vertebrates

there is a selection pressure on

their prey species to differ from

one another in appearance. The

effects of this selection pressure

will be greatest where predator-

-prey relationships are most com-

plex. The available evidence sup-

ports the hypothesis but experi-

mental results are needed to either

accept or reject it. The problem of

aspect diversity is worthy of fur-

ther study. The evolution of diver-

sity and of crypsis are both among

the important facets of our central

problem, the evolution of life.

It would be easier to document

ideas about tropical diversity if the

taxonomy, distribution and ecology

of tropical organisms were better

known but it will be many years

before a complete biological invent-

ory of the tropics is completed. It

is important in tropical biology to

study not only the animais and

plants themselves but the rela-

tionships between them on as

many functional and conceptual

leveis as possible.

Acknowledgments — I would like to

thank M. H. Robinson and M. Nau-

mann whose stimulating comments

helped me clarify my ideas. Drs. N.

Smith, W. J. Smith, and I. Rubinoff

kindly commented on the manuscript.

SUMMARY

The evolution of the diversity of

tropical life, that is so striking in

Amazônia, is a basic problem for

tropical biology today. The facet of

diversity most studied has been

species diversity. Tropical life is

also characterized by its variety of

forms and appearance, or aspect

diversity. To explain aspect diver-

sity I would suggest the hypotheses

that : A. Because learned character-

istics play an important role in

prey recognition by visually orient-

ed vertebrates there is a selection

pressure on their prey species to

differ from one another in appear-

ance. B. The effects of this selec-

tion pressure will be greatest where

predator-prey relationships are

most complex.

Evidence in the literature de-

monstrates that predators can in-

fluence the appearance of prey and

that at least some birds and mam-

mals in controlled experiments im-

prove with practice in their ability

to find cryptic prey. Studies on

Tits in Holland show that learning

can play a role in efficiency of prey

location in the wild and that prey

species are learned separately.

Volume 5 (Zoologia)

81

Because each time a predator

finds a particular prey it tempo-

rarily improves its ability to find

the next one of that type, there is

a selective advantage for each spe-

cies to look as different from all

others as possible so that a preda-

tor’s learning task is as difficult

as possible. This is selection for as-

pect diversity.

Examination of several samples

shows that sympatric insects oí

similar basic forms and using sim-

ilar types of camouflage still

differ in appearance. This is pre-

dicted by the hypothesis.

Among other possible effects of

predator learning on prey species

are increased variability in com-

mon species, maintenance of poly-

morphism and evolution of mono-

phagy in similar caterpillars, and

promotion of species diversity.

Tropical faunas have very com-

plex predator-prey relationships

and these lead to strong selection

pressures for differences among

cryptic animais and this helps ex-

plain the extreme evolution of as-

pect diversity in the tropics.

SUMÁRIO

A evolução da diversidade da

vida tropical, que é tão marcante

na Amazônia, é um problema bá-

sico para a biologia tropical de

hoje. A faceta mais estudada tem

sido a diversidade das espécies. A

vida nos trópicos é também carac-

terizada por sua variedade de for-

mas e aparência, ou diversidade de

aspecto. Para explicar a diversida-

de de aspecto, eu sugeriria as hipó-

teses: A) Como as características

aprendidas desempenham um im-

portante papel no reconhecimento

das presas pelos vertebrados orien-

tados pela visão, há uma pressão

de seleção em relação às suas pre-

sas, de modo a separar cada qual

pela aparência; B) Os efeitos desta

pressão de seleção serão maiores

onde as relações de predador-pre-

sa forem mais complexas.

Evidências na literatura demons-

tram que os predadores podem in-

fluenciar a aparência da presa e

que, pelo menos, algumas aves e

mamíferos em experimentos con-

trolados, melhoram, com a prática,

suas habilidades em encontrar a

presa camuflada. Estudos em sa-

guis, na Holanda, mostram que a

aprendizagem pode desempenhar

um papel na eficiência da locali-

zação da presa, na natureza, e que,

para cada presa, a aprendizagem é

feita separadamente.

Como cada vez que um predador

encontra uma determinada presa,

suas habilidades para achar outra

semelhante vão progressivamente

melhorando, há uma vantagem se-

letiva em que cada espécie possa

parecer tão diferente de tôdas as

outras o mais possível, isto é, para

dificultar cada vez mais o traba-

6 — 37 145

82

Atas do Simpósio sôbre a Biota Amazônica

lho de um predador em aprendiza-

gem. Isto é a seleção para a diver-

sidade de aspecto.

Exames de várias amostras dizem

que os insetos simpátricos de for-

mas básicas similares e usando se-

melhantes tipos de camuflagem,

ainda diferem na aparência. Isto

é uma hipótese.

Entre outros possíveis efeitos do

predador aprender nas espécies de

presas, estão, a variabilidade cres-

cente em espécies comuns, a ma-

nutenção de polimorfismo e evolu-

ção da monofagia em lagartas se-

melhantes e a promoção da diver-

sificação das espécies.

As faunas tropicais têm relações

predador-presa bastante comple-

xas e estas conduzem a fortes

pressões de seleção, pelas diferen-

ças entre os animais camuflados,

o que ajuda a explicar a extrema

evolução da diversidade de aspecto

nos trópicos.

LITERATURE CITED

Brower, J. V. Z., 1960, Experimental

studies of mimiery. IV. The reac-

tions of starlings to different pro-

portions of models and mimics.

Amer. Nat., 94 (877) : 271-292.

Brower, J. V. Z. & Brower, L. P., 1965,

Experimental studies of mimiery.

VIII. Further investigations of Ho-

neybees ( Apis mellifera ) and their

Dronefly mimics (Eristalis ssp.) .

Amer. Nat., 99 (906) : 173-187.

Brower, L. P., 1958, Bird predation and

foodplant specificity in closely re-

lated procryptic insects. Amer.

Nat., 92 (864): 183-187.

Cain, A. J. & Sheppard, P. M„ 1950, Se-

lection in the polymorphic land

snail Cepaea nemoralis. Heredity,

4: 275-294.

Cain, A. J. & Sheppard, P. M., 1954, Na-

tural selection in Cepaea. Genetics,

39: 89-116.

Cott, H. B., 1940, Adaptive coloration

in animais. 508 p. Methuen & Co.,

London.

Ford, E. B., 1964, Ecological genetics.

335 p. Methuen & Co., London.

Herrebout, W. M., Kuyten, P. J. &

Ruiter, L„ 1963, Observations on

colour patterns and behaviour of

caterpillars feeding on Scots pine.

Arch. Néerl. Zool., 15 (3) : 315-357.

Inger, R. & Marx, H., 1961, The food of

amphibians. Explor. Pare National

de VUpemba, fase. 64: 86 p.

Kettlewell, H. B. D., 1959, Las adapta-

ciones en los insectos brasilenos.

Endeavour, 18 (72) : 200-210.

Kettlewell, H. B. D., 1961, The pheno-

menon of industrial melanism in

Lepidoptera. Ann. Rev. Ent., 6:

245-262.

Klopfer, P. H. & MacArthtjr, R. H„

1960, Niche size and faunal diver-

sity. Amer. Nat., 94: 293-300.

Mook, J. H., Mook, L. J. & Heikens, H.

S„ 1960, Further evidence for the

role of “searching images” in the

hunting behaviour of titmice. Arch.

Néerl. Zool., 13: 448-465.

Paine, R. T., 1966, Food web complexity

and species diversity. Amer. Nat.,

100 (910) : 65-75.

Pianka, E. R., 1966, Latitudinal gra-

dients in species diversity: a review

of concepts. Amer. Nat., 100: 33-46.

Rilling, S„ Mittelstaedt, H. & Roeder,

K. D., 1959, Prey recognition in the

praying mantis. Behaviour, 14:

164-184.

Volume 5 (Zoologia)

83

Ruiter, L., 1952, Some experiments on

the camouflage of stick caterpil-

lars. Behaviour, 4: 222-232.

Sexton, O. J., 1960, Experimental stu-

dies of artificial batesian mimics.

Behaviour, 15: 244-252.

Sheppard, P. M., 1958, Natural selection

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Skutch, A. F., 1954, Life histories of

Central American birds. I. Families

Fringilidae, Thraupidae, Icteridae,

Parulidae and Coerebidae. Cooper

Ornithological Soc., Pacific Coast

Avifauna, No. 31, 448 p.

Skutch, A. F., 1960, Life histories of

Central American birds. II. Fami-

lies Vireonidae, Sylviidae, Turdidae,

Troglodytidae, Paridae, Corvidae,

Hirundinidae, and Tyrannidae.

Cooper Ornithological Soc., Pacific

Coast Avifauna, No. 34, 593 p.

Thorpe, w. H., 1956, Learning and ins-

tinct in animais. 493 p. Methuen &

Co., London.

Txnbergen, L., 1960, The natural con-

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tors influencing the intensity of

predation by songbirds. Arch.

Néerl. Zool, 13: 265-336.

Tinbergen, N., 1957, Defense by color.

Sei. Amer., 197 (4) : 48-54.

Uexküll, J. von, 1957, A stroll through

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Instinctive behavior. International

Universities Press, Inc., New York.

Atas do Simpósio sôbre a Biota Amazônica

Vol. 5 (Zoologia): 85-95 — 1967

PROBLEMS AND PROGRAMS IN AMAZONIAN ZOOLOGY

P. E. VANZOLINI

Departamento de Zoologia, São Paulo

Amazônia is at present the focus

of much deserved attention. On

one hand it as large area of prac-

tically continuous and little damag-

ed equatorial rain forest, placed

in the heart of a continent which

has just passed through a long pe-

riod of isolation, and its biota cer-

tainly contains clues to many cri-

ticai problems of evolutionary bio-

logy. On the other hand, its econ-

omic potentials are practically

undeveloped from the viewpoint of

exploitation of natural resources,

as well as of human occupation,

both of which are at present acute-

ly urgent matters. It indeed repre-

sents at the same time an im-

mensely valuable patrimony and a

heavy responsibility to the coun-

tries which hold sovereignty over it.

Anything that is done about

Amazônia has to be done quickly

and well. The problems are huge,

the retard in attacking them al-

ready considerable, and the viabi-

lity of adequate Solutions much

threatened by the inordinately in-

creasing pressure of the blind for-

ces of demographic expansion and

comercial interest.

It is the intention of this paper

to review some areas of zoological

research of immediate concern and

to submit approaches, theoretical

and practical, to their study.

ECOLOGY OF THE KEYSTONE

From whatever angie I look

at Amazonian problems, ecology

looms to me as the central issue

in the biological area. Efficient use

of natural resources obviously hin-

ges on possession of accurate au-

tecological and synecological data.

Human settlement, with the at-

tendant health and agricultural

problems, already known to be

rather difficult, depends heavily on

awareness and control of ecological

features. And the contributions of

the study of the Amazonian flora

and fauna to evolutionary biology,

the central theme of this paper,

will certainly spring mainly from

86

Atas áo Simpósio sôbre a Biota Amazônica

an understanding of the interrela-

tions between the living and the

physical environments, as well as

of the internai adjustments which

permit an extremely complex biota

to evolve and to maintain itself.

Adequate concentration of mate-

rial resources and manpower can

afford significant results in a short

time. However, it seems that there

is one deterrent to intensive ecolo-

gical studies in Amazônia, the ina-

dequate taxonomic knowledge of

the fauna, and possibly of the flora.

Thus, the first zoological aspect to

consider is the need of cataloguing

the fauna.

FAUNISTIC CATALOGUES

I mean by faunistic catalogues

not simple lists of species and

their places of occurrence, but data

originating from revisions of ample

scope, even if not necessarily in-

volving evolutionary and zoogeo-

graphical analyses.

The essential elements for this

type of work are, of course, large

primary collections from the area

of interest, and access to the lite-

rature and to reference collections,

especially those containing types.

The assembling of large collec-

tions, both conceptually and as a

practical matter, will be separately

discussed below. The other mate-

rial elements are nowadays perfect-

ly accessible to anyone, even to

zoologists working in South Ame-

rica. Some local institutions have

progressed to the point where a

fair share of the identifications can

be done at a first approach. Inter-

national cooperation and modern

means of communication have

made the key libraries and collec-

tions easy to consult. In fact,

enough experience has accumulat-

ed to indicate that there are no

outstanding difficulties in the way

of major revisions.

However, if material resources

are not a hindrance to intensive

taxonomic studies of South Americ-

an animais, one must concede that

there is a definite lack of trained

personnel. It is therefore an ur-

gent task to increase the number

of competent taxonomists.

I say “South American”, and not

“Amazonian”, advisedly. I believe

that any systematic revision with

pretensions to a degree of perma-

nence has to take into account at

least a major zoo-geographical

unit, in the present case all of the

Guiano-Brasilian subregion. Ama-

zônia in itself is not a unit from

the viewpoint of evolutionary tax-

onomy.

This concept has two important

practical consequences. First,

training for Amazonian taxonomic

research should always include an

adequate measure of field work in

the Atlantic forest and in the Cen-

Volume 5 (Zoologia)

87

trai Brasilian cerrados, and possi-

bly in the northeastern caatingas,

in order to give the zoologist

enough ecological competence to

interpret his distributional data

and patterns of differentiation. In

all other respects, formation of

zoologists can perfectly follow tra-

ditional lines : broad theoretical

background, sound craftsmanship,

and a good infusion of practical

ecology, given in the terrain. More

will be said of this when dealing

with intensive collecting efforts.

Second, Amazonian collections

by themselves do not mean much.

They must be part of a large en-

semble, encompassing the Guiano-

-Brasilian, subregion for imme-

diate taxonomic purposes, and as

much more as necessary for com-

parative work.

EVOLUTIONARY TAXONOMY

It was said above that mono-

graphic work with faunistic inten-

tions need not forcibly go into evo-

lutionary or zoogeographical mat-

ters. Of course this does not mean

that it should not; on the contrary,

as soon as the available informa-

tion warrants it, the man who laid

the groundwork is a fool not to

reap as much as he possibly can of

the theoretical fruits.

South American zoology affords

many good opportunities for such

work. On a primary levei are accu-

rate descriptions of differentiation

patterns, within any significant

taxon. The accumulation of such

works leads to the recognition

of regularities and singularities

which, correlated with contempo-

rary and historical morphoclima-

tic patterns, contribute new infor-

mation and independent checks to

theoretical work done on the tem-

perate and Old World tropical fau-

nas.

Let us consider some relevant

issues.

Morphoclimatic domains

The Neotropical region can be

reasonably divided into four subre-

gions: West Indian, Centro- Ameri-

can, Andino-Patagonian and Guia-

no-Brasilian, the last named being

the core of Neotropica.

Within the Guiano-Brasilian su-

bregion one can consider four

major morphoclimatic domains: 1

the Amazonian (and Guianan)

hylaea, the Atlantic forest, the

Central-Brasilian cerrados and the

caatingas of northeastern Brasil.

The presence and role of smaller

areas of other types, and of fácies

and geographical subdivisions of

the four domains, will become

eventually (one hopes shortly) sig-

1 Thanks are due to Aziz Nacib

Ab’Saber, Universidade de S. Paulo, for

his unfailing help in geographical mat-

ters, both in the library and in the

field. The present concept of Brasilian

morpho-climatic domains is his.

88

Atas do Simpósio sobre a Biota Amazônica

nificant to zoology. It is probable,

however, that the main evolution-

ary and distributional features of

the fauna will be shown to depend

on the static and dynamic rela-

tionships among the four major do-

mains, with a contribution from

marginal areas. At any rate, the

information now available permits

only the consideration of the four

domains.

Their present distribution is of

paramount interest. The two rain

forest areas are separated by the

broad belt of open formations (cer-

rados and caatingas), which runs

from SW to NE. There is only a

slight latitudinal overlap between

the Southern edge of the hylaea (a

festooned line broadly wavering

about the 10th parallel) and the

narrow northern coastal wedge of

the Atlantic forest in Pernambuco

and Paraíba (approximately 7o).

The core of the cerrados lies to

the SW of the core of the caatin-

gas. There is ample contact be-

tween them and, in places deter-

mined by topography, a strong

measure of interdigitation. Latitu-

dinal overlap occurs between the

4th and the 16th parallels.

The rivers which run from the

core of the cerrado into the Ama-

zonas are bordered by gallery for-

ests from the very head waters.

The width of these ciliary forma-

tions is, within the cerrado core,

closely proportional to the width

of the water course. In northern

Mato Grosso they start spreading

until the galleries of adjacent ri-

vers fuse and one finds oneself in

the hylaea. The relatively recent

advant of extensive air travei in

the interior has shown that signi-

ficant interfluvial cerrados extend

much farther north than river ex-

ploration had previously led to be-

lieve. The eastern portion of the

hylaea-cerrado contact, in Mara-

nhão, follows broadly an interflu-

vium, and may be complicated by

the presence (also probable else-

where) , of drier subtypes of forest.

In spite of deficiencies of map-

ping and floristic studies, it can be

said that each one of the four do-

mains contains enclaves of forma-

tions characteristically belonging,

at least physiognomically, to one

or two of the others. Mutual out-

liers of cerrado and caatinga are

known to exist all along the con-

tact. Cerrado or cerrado-like areas,

some quite extensive, occur in

Amazônia and within the Atlantic

forest. Isolated stands of forest,

smaller or larger, are not unfre-

quent in the cerrado. In the caatin-

ga, they also exist (“brejos”) and

are clearly determined by local oro-

graphical conditions.

Also pertinent to the present ar-

gument are two areas marginal to

Amazônia: the Andes of Ecuador,

northern Peru and Colombia, and

Volume 5 (Zoologia)

89

the open formations (“savannas”)

of Venezuela and the Guianas.

The forest climbs up the Andes

quite a ways before petering out,

and, especially in Ecuador, deep

river valleys are forested right up

to the middle of the cordillera.

No adequate botanical compari-

son is available to a zoologist of

the open formations north and

south of the hylaea, but the mere

fact that they are open is very im-

portant in itself.

Major ecological preferences of the

fauna

Many elements of the South

American fauna have an extremely

broad ecological spectrum, occurr-

ing in practically all environ-

ments. On the other extreme, there

are forms restricted to special ha-

bitats, such as small geographic

areas or specific food plants. It is

also safe to assume that a number

of open-formation species will be

found to be, preferrably or exclu-

sively, adapted to either cerrado or

caatinga, independently or clima-

tic zonation.

However, a sizable proportion of

the fauna shows an intermediate

degree of ecological specialization,

being adapted to either open or

closed formations, but not obeying

other restrietions, except broad la-

titudinal (or, better, temperature)

zonation. This segment of the fau-

na is, at present, the most reward-

ing for evolutionary studies, espe-

cially with regard to general pat-

terns of speciation and to the phe-

nomena related to the rise and per-

manence of complex biotas and,

consequently, of dominant faunas.

Immediate habitats

Among the implicit assumptions

of zoologists about the hylaea,

ranks foremost a belief in its eco-

logical homogeneity, or at best, in

the traditional tripartite division:

“terra firme”, “várzea” and “iga-

pó” — dry-land, back-swamp and

permanently-flooded forest. But

any amount of purposeful collect-

ing of small vertebrates will show

immediately the importance of va-

riations in the immediate habitat,

perhaps too gross to be called eco-

logical niches. In my experience,

the most important is the relative

amount of light and shade, and

consequently the distribution of

small open spaces. There is a defi-

nite proportion of ecotonal ani-

mais, found only on the first me-

ters of the forest; there also is a

fauna of the small clearings and

trails, in several instances cutting

across the major ecological divi-

sions. In fact, some animais broad-

ly distributed through the heart

of the valley do not belong directly

to the dark humid forest.

An awareness of this aspect is

fundamental to work on geograph-

90

Atas do Simpósio sôbre a Biota Amazônica

ical differentiation and on ori-

gins and pathways of faunistic

units.

History of the fauna

The study of fóssil mammals has

led to the consideration of three

f aunai strata in South America:

one autochthonous element, resi-

dent since at least the Upper Me-

sozoic; one recent (Upper Plioce-

ne) immigrant from North and

Central America; and an interme-

diate, much smaller group of Mid-

dle T e r t i a r y “island-hoppers” .

I would add a fourth category, also

small, for those forms which were

rafted across either the Atlantic or

the Pacific2.

Paleontological and zoogeograph-

ical data on other groups tend

to give general validity to this con-

cept.

The relative weight of the strata

varies from taxon to taxon. For

instance, recent immigrant mam-

mals constitute a significant por-

tion of the mastis. On the contrary,

South American fishes, chelonians,

and crocodylians, are strongly en-

trenched autochthonous groups,

having admitted practically no in-

vaders. Snakes and lizards are in-

termediate in this respect.

The area of first encounter of

northern immigrants and old re-

3 For a discussion of fauiial strata

and related matters see Vanzolini &

Guimarães (1955) and Vanzolini (1964).

sidents in obviously the northwest-

ern corner of the continent. In

fact, many northern groups have

their only foothold in South Ame-

rica in the trans-Andean belt from

northern Peru to Venezuela. Simp-

son has called this a “filter area”,

but I would prefer a name “north-

western arch”, with no causal

connotations, as among the forms

presently restricted to the belt are

some which have been, in the past,

widespread in the continent.

Except for the Anãean highway,

all routes of immigration past the

northwestern arch must cross

Amazônia. It is reasonable to ex-

pect to find there movements in

progress and the traces of past phe-

nomena.

Paleoclimatology

Until quite recently, paleoclim-

ates did not enter much into the

cogitations of zoologists working

on South America. There was, on

one hand, very little information;

on the other, an implicit belief in

the slowness and even in a rela-

tive unimportance of climate

changes, especially in the inter-

tropical belt.

The outlook has changed com-

pletely in recent years. Evidence

from many sides has been accumu-

lating very fast, all of it pointing

to extremely drastic and quite re-

cent climatic changes. In fact, the

information presently being ac-

Volume 5 (Zoologia)

91

quired comes from so many fields

and grows so fast that it is very

difficult at present to prepare a

balanced abstract. Orthodox pollen

studies, analyses of archeological

sites geomorphological studies of

mountain profiles and of sand

beaches, are some of the sources of

paleoclimatic data presently in in-

tensive progress.

These studies make it perfectly

clear that, in the last few thousand

years, some areas such as, for

example, western Paraná and large

tracts of the Serra do Mar (the

present stronghold of the Atlantic

forest) have been at least devoid of

closed formations and in all pro-

bability actually semi-arid. In the

Serra do Mar there is also evidence

of the persistence of refuges —

certainly similar to present oro-

graphic “brejos” in the caatingas

— where the forest found condi-

tions to survive during the dry

epochs.

It is also clear that, during the

upper Tertiary and Quaternary,

there were several such cycles.

The simple awareness of these

data has a great impact on our

zcological thinking. For example,

a great deal of “ecological specia-

tion” becomes unnecessary to ex-

plain the large numbers of sympa-

tric closely related species observed

in some presently homogenous ex-

tensive areas.

Solution of actual problems is

still at best possible only for some

special cases.3 Evidence of one

type of climate for a certain place

at a certain time cannot be ex-

tended to broad areas; it suffices

to remember that hylaea, cerrados

and caatingas are found today on

the same latitudes and not far

apart.

As things stand, we zoologists are

in impatient need of much detail-

ed paleoclimatic information, de-

rived from an extensive geograph-

ical coverage and also from infe-

rential studies of dynamic climato-

logy.

A program for taxonomic studies

The preceding considerations

afford a first setting for a taxonom-

ic program. First of all, it must

be remembered that, unless the

group is naturally so restricted,

taxonomic work cannot be limited

to Amazônia. At least the Guiano-

-Brasilian subregion has to be cov-

ered, since the patterns of diffe-

rentiation should be at the very

least correlated to the contrasts

between the morphoclimatic do-

mains.

a A very favourable case is that of

the iguanid lizard Liolaemus lutzae,

whose separation from the parent

species can be confidently dated (Van-

zolini, unpublished) . A more conjec-

tural analysis has been made of the

iguanid lizard Anolis transversalis

(Williams & Vanzolini, 1966).

92

Atas do Simpósio sôbre a Biota Amazônica

Especially important are the dis-

junct groups that extend into the

Atlantic forest but not into the

open formations. They offer valu-

able inshights on rates of differen-

tiation and on the effects of clima-

tic zonation.* * 4

Taxa which occur both in the

open and closed formations must

be studied from the viewppint of

what was called above the “imme-

diate habitat”. Are they umbro-

phils who frequent the gallery for-

ests of the cerrado or heliophils

who enter the forest along clear-

ings and trails? Or yet really

adapted to both environments, or

in process of adaptation? Both the

methodology and conclusions of

the analysis depend on this type of

information.

Special attention must be given,

of course, to the fauna of enclaves

and to comparisons of the open

formations which lie to the north

and south of the hylea.5

In this way the progress of ca-

taloguing will carry along both

evolutionary information of imme-

* Some important cases of disjunct

distributions are listed in Vanzolini,

1962.

5 The faunal dissimilarity between

the open formations lying north and

south of the hylaea is at times strik-

ing. Among others, the several species

of deer, the rhea, the open-country ti-

namids, etc., which range very widely

in Brasil, up to Southern Pará, are not

present in the Guianas. In contrast,

the rattlesnake, Crotalus durissus, is

found on both sides and in enclaves.

diate usage and an accumulation

of data for eventual syntheses of

the historical processes through

which the Amazonian — and, in

general, South American — fauna

passed.

COMPLEX BIOTAS

A major field of modern zoologic-

al research is the origin and per-

manence of complex biotas, to

which is closely linked the problem

of the rise of dominant faunas.

The usual explanation seems to

me tautological. It is frequently

said that the tropical regions are

more favourable to life. The expres-

sion “more favourable” cannot

mean “permitting a larger quan-

tity of life”; I do not believe there

is more living matter in the hylaea

than in the redwood forest. If

“more favourable” means “permit-

ting larger diversity”, there we are

back at the beginning.

Amazonian research permits se-

veral lines of attack. To begin,

quantitative surveys of the mor-

phoclimatic domains are needed to

afford a better concept of what

is meant by the loose but current

term “tropical”.

Oriented autecological and syne-

cological analyses are necessary to

understand the beginning of diver-

sification. A biota already diversi-

fied offers many opportunities for

further diversification, as subtler

differences between ecological ni-

Volume 5 (Zoologia)

93

ches become possible and, obvious-

ly, are used. Once the process is

initiated, one can expect a positive

feedback to be installed; the situa-

tion of the temperate biotas may

be explained by a poorer beginning

and perhaps a negative feedback.

Considering the large amounts

of living matter produced under

temperate climates, I’m led to

think that the initial push to the

building of a diversified biota is

the longer reproductive season

available. The first, easiest multi-

plication of ecological niches would

be the apportioning of the oppor-

tunities for reproduction. I believe

research along these lines to be

extremely promising.

On another tack, a diversified

biota is a good subject for investi-

gation on the concept of ecological

niche, especially with regard to

Gause’s principie. The meaning of

small diferences in preferred food,

for instance, can be much better

investigated in the presence of a

wide range of subtle choices.

Finally, the ways in which living

space is shared and how the ill

effects of crowding on communica-

tion are avoided are best studied

in the taxonomically wealthy rain

forest.

COLLECTIONS

We come now to a most import-

ant practical problem — the as-

sembling of primary collections

permitting intensive taxonomic

work on the groups significantly

represented in Amazônia. The fol-

lowing ideas are based on my per-

sonal experience of Brasilian Ama-

zônia.

The making

The first aspect to be considered

is logistics. Present and foreseeable

conditions in Brasil are such that

any sustained collecting effort

must be made along the navigable

rivers. Furthermore, it must count

on its own means of transporta-

tion. The small (10-12 ton) “mo-

tores” of the region are very con-

venient, as they are well fitted to

local conditions and reliable crews

easily available. I believe an en-

semble of a regular diesel-driven

boat and a towed barge (for labo-

ratory and living quarters) , plus a

complement of canoes, to be ideal.

Explora tion above the falis, es-

pecially of the eastern tributaries

of the left bank, is a more difficult

proposition, which can only be solv-

ed with the help of itinerant trad-

ers, who have specialized equip-

ment and personnel.

Work on the interfluvia is easy

in the lowlands, where large canoes

can do any amount of penetration.

In the northern and Southern

uplands one must have recourse to

the Brasilian Air Force and to local

expeditions based on the river

boats.

94

Atas do Simpósio sôbre a Biota Amazônica

The flotilla system has another

advantage. Regular living and

working quarters for 8 or 10 na-

turalists permit the assembling of

interdisciplinary crews. It is no

news that this type of team greatly

enriches the work, but we must

stress that in the case of Amazônia

this is doubly important.

In fact, from what has been dis-

cussed above it should have be-

come clear that simple locality re-

cords are not sufficient for good

revisionary work. Notes on the

exact habitat and on the general

features of the area are desirable.

We totally lack texts which would

prepare a taxonomist to fully un-

derstand the landscape, and every

one of us has to struggle for him-

self to acquire some acquaintance

with the relevant ecological fea-

tures. A team including ecologists

and botanists is capable of gather-

ing much better data than the

isolated zoologist, and joint work

will do much to educate everybody

in fields outside his own.

A practical experiment along

these lines is under way. A group

of institution, Brasilian and fo-

reign, is undertaking a program of

ichthyological research on the mid-

dle Amazonas. For this purpose,

two boats (tug and lab barge) are

presently being built in Pará. It is

intended that, between collecting

trips and after the program is com-

pleted, these boats will be placed

on full time Service along the lines

advocated above. They will be

made available to interdisciplinary

and inter-institutional groups for

intensive collecting work. Since the

costs of building and outfitting

will be absorbed by the initial pro-

ject, subsequent field work will be

relatively inexpensive.

This program will have the

added advantage of allowing for

the training of students. It is our

belief that every prospective Brasil-

ian zoologist, whatever his intend-

ed field of work, should spend a

period of practical training in

Amazônia. This will be obviously

true for many people in other

countries. A permanent and inex-

pensive expedition will bring this

goal closer to attainment.

Placement of collections

Custom, ethics and reason deter-

mine that as representative as pos-

sible a part of every collection re-

main in the hands of the zoologist

who identified or otherwise stu-

died it.

The main body of a collection

however, as well as all possible

additions to it, should remain in

one institution where the most ef-

ficient use can be made of it. In

the case of Amazônia, which is a

focus of interest to the whole West-

ern world, this means that the

hylaean countries should be pre-

pared to house and care for large

Volume 5 (Zoologia)

95

collections and to keep them avail-

able to qualified scientists.

Fortunately this is already pos-

sible. Several Latin American ins-

titutions are in a position to share

in this task. Furthermore, institu-

tional and scientific relations are

extremely cordial and one can ex-

pect no trouble to formal or infor-

mal agreements on where to keep

the bulk of the collections of

individual taxa. A consideration

of existing collections (especially

those complementary to Amazo-

nian ones) , tradition and other cir-

cumstances should make it easier

still.

One special type of collection

should also be contemplated: sy-

noptic regional collections intend-

ed for the use of ecologists, phy-

siologists, pathologists and other

people who work on the fauna for

non-taxonomic purposes. The col-

lections, made with the cooperation

of all systematists, should be

housed in institutions seated in

the region.

CONCLUSION

This is a brief and cursory

sketch which certainly reveals too

much of my taxonomic background

and interests. I hope, however, that

it can serve as a preliminary frame

of reference for a discussion of how

to do something, quickly and well,

about Amazonian zoology.

LITERATURE CITED

Bigarella, J. J. & Andrade, G. O., 1965,

Contribution to the study of the

Brazilian Quaternary. Geol. Soc.

Amer. Special Paper, 84: 433-451.

Bigarella, J. J., Marques F.°, P. L. &

Ab’Saber, a. N„ 1961, Ocorrências

de pedimentos remanescentes nas

fraldas da Serra do Iquererim (Ga-

ruva, SC) . Boi. Paranaense Geogr.,

4/5: 82-93.

VAN DER HAMMEN, T. & GONZALES, E.,

1965, A Late Glacial Holocene pol-

len diagram from Cienaga dei Vi-

sitador (Depto. Boyacá, Colombia! .

Leidse Geol. Med., 32: 193-201.

Lamlng, A. & Emperaire, J., 1959, A ja-

zida José Vieira, um sítio guarani

e pré-cerâmico do interior do Pa-

raná. Publ. Cons. Pesquisas Univ.

Paraná, Curitiba, Arqueologia, 1:

143 pp.

Tricart, J., 1958, Division morphocli-

matique du Brésil Atlantique Cen-

tral. Rev. Geomorph. Dynam., 9:

1-2.

Vanzolini, P. E., 1962, Problemas fau-

nísticos do cerrado, in Simpósio

sôbre o cerrado, pp. 307-320. S.

Paulo: Editora da Universidade de

S. Paulo.

Vanzolini, P. E., 1964, Lagartos brasi-

leiros da família Gekkonidae: Sis-

temática, evolução e geografia (Sau-

ria) . 217 pp. Tese apresentada à

Faculdade de Filosofia, Ciências e

Letras da Universidade de S. Paulo.

Vanzolini, P. E. & Guimarães, L. R„

1955, Lice and the history of South

American Land mammals. Rev.

Brasil. Ent., 3: 13-46.

Williams, e. E. & Vanzolini, P. E„

1966, Studies on South American

Anoles. Anolis transversalis A. Du-

méril. Papeis Av. Dep. Zool. S.

Paulo, 19: 197-204.

Atas do Simpósio sôbre a Biota Amazônica

Vol. 5 (Zoologia): 97-101 — 1967

DIE BODENFAUNA DES NEOTROPISCHEN REGENWALDES

LUDWIG BECK

Institut für Allegemeine Zoologie der Ruhr-Universitàt, Bochum und Instituto

Nacional de Pesquisas da Amazônia, Manaus

Der Regenwald Amazoniens, der

den grõpten Teil der neotropis-

chen Region bedeckt, ist auffallend

arm an Groptieren, besonders Sáu-

getieren, die in grofSer Zahl die pa-

làotropische Region bevõlkern. Die-

ser Armut steht jedoch ein enormer

Reichtum an Arthropoden gegenii-

ber und somit auch eine mannig-

faltige Bcdenfauna, die gerade im

Regenwald im wesentlichen aus

Arthropoden besteht.

Einer der ersten und auffallen-

dsten Merkmale ist die Allgegen-

wart zweier Insektengruppen, der

Ameisen und Termiten. Williams

(1941) gibt den Anteil der Amei-

sen an der Bodenfauna eines Re-

genwaldes in Panama mit etwa 1/ 4

der Individuendichte an, und wir

kõnnen diesen Anteil ebenso für

den Regenwald des Amazonasge-

bietes annehmen. Noch deutlicher

wird die beherrschende Stellung

der Ameisen, zieht man die Kõr-

pergrõpe oder Biomasse in Betra-

cht. Ahnliches gilt für die Termi-

ten, die in groper Individuenzahl

alie “Schichten” des Waldes besie-

deln. Ihr Prozentanteil an der Bo-

denfauna ist infolge ihrer Lebens-

weise kaum exakt zu ermitteln;

doch tíürfte er dem der Ameisen

kaum nachstehen und zusammen

mit diesen stellen die Termiten si-

cher über 3/4 der Biomasse der Bo-

denfauna.

Neben Ameisen und Termiten

lebt eine Fülle von Tiergruppen im

Boden, von denen keine beherrs-

chend hervortritt. Man kann sie

recht gut quantitativ erfassen mit

der Berlese — Methode: In einen

gropen Trichter wird oben ein

Drahtnetz eingesetzt, worauf die

Bodenprobe gebreitet wird. Darü-

ber befestigt man eine Glühbirne,

die die Probe langsam von oben

her erwarmt und austrocknet. Die

Tiere weichen vor Licht, Wárme

und Trockenheit nach unten aus

und fallen durch den Trichter in

ein darunter gestelltes Sammel-

glas.

7 — 37 145

98

Atas do Simpósio sôbre a Biota Amazônica

Von den damit erfajlten kleine-

ren Boden-Arthropoden überwie-

gen zahlenmâpig die Milben, im

wesentlichen Oribatiden. Etwa 100

Arten sind von ihnen bisher aus

dem Regenwald bekannt und ihre

gesamte Artenzahl dürfte ein Meh-

rfaches ausmachen. Wir kennen

einige, hier endemische Gattun-

gen: Epactozetes, Schalleria, Dino-

zetes. Viele circumtropisch verbrei-

tete Gattungen wie Rostrozetes,

Heterobelba Dampfiella, Rhyncbo-

ribates, Acronothrus und Lamello-

bates sind hier mit endemischen

Arten vertreten. Nur 4 Oribatiden-

Arten des neotropischen Regen-

waldes wurden auch au|3erhalb

dieser Region gefunden (Beck,. . .

1963).

Nach den Milben stellen die Col-

lembolen die náchstháufige Boden-

tiergruppe; sie ist durch zahlreiche

Arbeiten besonders von Arlé, Bon-

net, Denis und Winter gut bekan-

nt. Auch hier kennen wir mehrere

für den Regenwald typische Gat-

tungen, so die Gattungen der Un-

terfamilie Paronellinae, Dicrano-

centrus und Discocyrtus (Winter,

1963).

Mit gro(3er Regelmãpigkeit fin-

den wir verschiedene Arten von Di-

plopoden in der Bodenfauna, wei-

terhin Kàfer (Staphyliniden, Pti-

liiden) , verschiedene Insektenlar-

ven (besonders von Coleopteren

und Dipteren), Asseln und Scha-

ben. Hiermit habben wir nur die

wichtigsten Bodentiergruppen ge-

nannt, neben denen man noch zah-

lreiche andere oftmals antrifft:

Symphylen, Pauropoden, Chilopo-

den, Spinnen, Pseudoskorpione,

Dipluren, Proturen, Thysanuren

und Wanzen.

Hingegen treten die in Bõden ge-

màfiigter Klimazonen in groper

Zahl vorkommenden verschiede-

nen Gruppen der “Würmer” wie

Nematoden und Oligochaeten zur-

ück. Zwar finden wir kleine Nema-

toden und Enchytraeen in der

Laubstreu, doch fehlen gerade die

sonst haufigen und wichtigen eue-

daphischen Formen im Mineral-

boden. Regenwürmer trift man nur

in geringer Zahl an. Doch wàhrend

Regenwürmer sonst tiefe Gánge im

Boden graben, organisches und

anorganisches Material vermis-

chen und in ihrem Darmtrakt die

wichtigen Tonhumuskomplexe bil-

den, dringen sie hier nicht oder nur

flach in den Boden ein, vielfach

kriechen sie nur in und unter der

Laubstreu umher.

Will man die Bedeutung der ein-

zelnen Tiergruppen für den Boden,

insbesondere für die Humuspro-

duktion, erschliepen, ist die Háu-

figkeit des Auftretens einer Tier-

gruppe und ihre Individuendichte

nur eines von mehreren Kriterien.

Man mu(3 darüber hinaus die Bio-

masse, wenn mõglich die Stoffwe-

chsel-Intensitàt und vor aliem Le-

Volume 5 ( Zoologia )

99

bensweise und Nahrungsaufnahme

berücksichtigen.

Für die Humusbildung ist der

Abbau des pflanzlichen Bestande-

sabfalls (Laub und Holz) durch

Bodentiere ausschlaggebend. Hier

spielen die Termiten die wichtigste

Rolle. Sie treten in ungeheuren In-

dividuenzahlen auf und verarbei-

ten gewaltige Mengen abgestorbe-

nen Holzes. Kein Baumrest, der

nicht von ihren Gàngen durchzo-

gen und ausgehõhlt wàre! Aller-

dings konzentrieren sie das verar-

beitete Material in ihren Bauten,

soda|3 ihm auf die Flàche bezogen

wesentlich geringere Bedeutung

zukommt ais der Menge nach

(Schaller, 1936).

An zweiter Stelle folgen, ihrer

Bedeutung für die Humusbildung

nach, die Diplopoden. Sie kommen

sehr háufig in der Laubstreu vor

und bei ihrer relativ grojlen Bio-

masse bedeuten etwa 15-20 Diplo-

poden pro 500 cm;! Substrat einen

wichtigen Faktor beim Abbau des

Pflanzenmaterials. Sie fressen auch

wening zersetztes Holz und sind

mit den Termiten ais Primár-Gro-

bzersetzer anzusehen.

Nach Termiten und Diplopoden

folgen in der Wichtigkeit bereits

die an Kõrpergrõjle sehr' viel klei-

neren Oribatiden, deren Individu-

endichte háufig zwischen 400 und

800 Tiere pro 500 cm:! Substrat be-

tràgt. Auch sie beteiligen sich nach

Schuster (1956) ais Primár-Grob-

zersetzer an dem Abbau der Laubs-

treu. Allerdings fressen Oribatiden

auch háufig Pilzhyphen, die sich

sehr schnell an abgestorbenem

Planzenmaterial entwickeln, und

es bedarf erst einer genauen Unter-

suchung, wieweit sich ihre Ernáh-

rung auf sekundáre Nahrungsstof-

fe der Laubstreu verlagert.

Das gilt auch für die Collembo-

len, die im Regenwald vermutlich

noch stárker auf sekundáre Nah-

rungsstoffe wie Pilzhyphen über-

gehen. Zwischen 250 und 500 Indi-

viduen leben durchschnittlich i n

500 cm3 Streu und beteiligen sich

zu einem gropen Teil am Abbau des

Fallaubs, das bereits einen gewis-

sen Zersetzungsgrad erreicht hat

(Schaller, 1950).

E i n i g e produktionsbiologische

Bedeutung kommt ferner den háü-

figen, aber meist nicht sehr zahl-

reichen Asseln ais Verzehrern von

F a 1 1 a u b und Holzresten zu;

man findet durchschnittlich 10-20

Exemplare in 500 cm:! Substrat.

Im Regenwald konkurrieren Pil-

ze und Bakterien in erheblichem

Umfang mit den genannten Tier-

gruppen bei der Zersetzung des

pflanzlichen Bestandesabfalls. Der

Befall durch Pilze vollzieht sich so

schnell, da|3 sich auch bei reinen

Makrophytenfressern, beispielswei-

se unter den Oribatiden, meist

Stücke von Pilzhyphen im Ver-

dauungskanal finden. Schlie|31ich

dürfte das enorme Bakterien-Wa-

100

Atas do Simpósio sôbre a Biota Amazônica

chstum bei optimalen Temperatur-

u n d Feuchtigkeits-Bedingungen

die Ursache für das fast võllige

Fehlen einer Humusschicht im Bo-

den sein. Durch Termiten, Diplopo-

den, Oribatiden, Collembolen, As-

seln und andere Tiere wird zwar

eine betráchtliche Menge Humus

in Losungsballen geliefert, die aber

sehr schnell dem Abbau in mine-

ralische Bestandteile anheimfallen.

Hier macht sich das Fehlen der

durch Regenwürmer gebildeten To-

nhumuskomplexe im Boden be-

merkbar. Die produktionsbiologis-

che Bedeutung der Bodentiere be-

darf aber noch eingehender Unter-

suchungen.

Ein überblick über die Bodenfau-

na tíieses Gebietes wáre unvolls-

tàndig, würden wir nur die boden-

biologisch bedeutenden Tiergrup-

pen erwàhnen. Es gehõren zahlrei-

che weitere Gruppen dazu, deren

Verhalten und Lebensweise für den

Zcologin sehr interessant sind;

sind es doch vielfach “altertümli-

che” Tiergruppen, deren Bauplan

sich seit Jahrmillionen unverán-

dert erhalten hat, und denen nur

der gleichmapig günstige Lebens-

raum ces tropischen Regenwaldes

das Überleben ermõglicht hat. Zu

ihnen gehõren mehrere Arachni-

den-Gruppen wie Amblypygi, Uro-

pygi, Palpigradi, und Ricinulei. Sie

leben alie in der Laubstreu ais

“Bodentiere” oder in bodennahen

Hõhlen und Lõchern am Fuji der

Baumriesen. Hier müssen wir auch

die Onychophoren erwàhnen, de-

ren Bauplan sie gleichsam ais

Bindeglied zwischen Anneliden und

Arthropoden ausweist; auch sie

finden wir regelmà|3ig in der Stre-

uschicht. Unter den Insekten ge-

hõren die Zorapteren und Embi-

iden hierher. Schlie|31ich sind noch

einige Tiere bemerkenswert, denen

das Regenwaldklima erlaubt hat,

das Wasser ais den ihnen “anges-

tammten” Lebensraum zu verlas-

sen und ais Landitiere zu leben:

So die Geoplanarien und einige

Crustaceen (Brachyuren).

So bietet uns die Bodenfauna des

neotropischen Regenwaldes ein

buntes, ungeheuer reichhaltiges

Bild, dessen Komposition uns nach

und nach bekannt ist, dessen tau-

send Einzelheiten aber noch der

Beschreibung und Erklãrung har-

ren.

Prof. Dr. F. Schaller und ich

versuchen zur Zeit, einige der an-

geschnittenen Probleme zu lõsen.

Mit Unterstützung der Deutschen

Forschungsgemeinschaft und ais

Gaste des Instituto Nacional de

Pesquisas da Amazônia in Manaus

und seines Direktors, Dr. Djalma

Batista, dem wir zu gropem Dank

verpflichtet sind, arbeiten wir im

einzelnen der Gõsung folgender

Fragen :

1. Qualitative und quantitative

Zusammensetzung der Bodenfauna

in verschiedenen Gebieten der Sch-

Volume 5 (Zoologia)

101

warz - und Weipwasserflüsse durch

Entnahme quantitativer Proben

nach der Berlese-Methode.

2. Ernáhrung und Nahrungsve-

rarbeitung einzelner Bodentier-

gruppen diirch Fütterungsversu-

che und Untersuchung des Darmi-

nhalts.

3 . Stoffwechsel - Intensitàt von

Bodentieren durch Messung des

Sauerstoffverbrauchs.

4. Biologie einiger seltener und

wenig bekannter Bodentiere.

ZUSAMMENFASSUNG

In der vorliegenden Arbeit wird

ein Überblick über die derzeit be-

kannte qualitative und quantita-

tive Zusammensetzung der Boden-

fauna des neotropischen Regenwal-

des gegeben. Die produktionsbiolo-

gische Bedeutung der einzelnen

Bodentiergruppen wird erõrtert

und unser Arbeitsprogramm weite-

rer Erforschung der Bodenfauna

in Amazonien skizziert.

SUMÁRIO

Êste trabalho contém um resumo

do conhecimento da composição

qualitativa e quantitativa da fauna

do solo da mata pluvial neotropi-

cal. Discute-se a importância dos

diferentes grupos de animais do

solo para a produção biológica e

faz-se um esboço de nosso projeto

de pesquisas sôbre a fauna do solo

na Amazônia.

LITERATUR

Beck, L., 1963, Zur õkologie und Taxio-

nomie der neotropischen Boden-

tiere I. Zur Oribatiden-Fauna

Perus. Zool. Jb., Syst., 90: 299-392.

Schaller, F., 1950, Biologische Beobach-

tungen an humusbildenden Bo-

dentieren. Zool. Jb., Syst., 78: 506-

-525.

Schaller, F., 1963, Bodenzoologische

Untersuchungen in Südamerika

und Afrika. Forsch. Fortschritte,

37: 100-104, 134-137.

Schuster, R., 1956, Der Anteil der Ori-

batiden an der Zersetzungsvor-

gángen im Boden. Z. Morph. okol.

Tiere, 45: 1-33.

Williams, E„ 1941, An ecological study

of the floor fauna of the Panama

rain forest. Buli. Chie. Sei. Acad.,

6: 63-124.

Winter, C., 1963, Zur õkologie und Ta-

xionomie der neotropischen Boden-

tiere II. Zur Collembolen-Fauna

Perus. Zool. Jb., Syst., 90: 393-520.

Atas do Simpósio sôbre a Biota Amazônica

Vol. 5 (Zoologia): 103-109 — 1967

NOTAS SÔBRE A DISTRIBUIÇÃO DE “BUFO GRANULOSUS”

SPIX , 1824 NA AMAZÔNIA E DESCRICAO DE UMA

SUBESPÉCIE NOVA (Amphibia, Bufonidae)

WERNER C. A. BOKERMANN

Departamento de Zoologia, São Paulo

(Com 8 figuras no texto)

Em trabalho recente, Gallardo

(1965) fêz um estudo das diversas

subespécies de Bufo granulosus ter-

minando por dividir a espécie em

14 subespécies.

Segundo as conclusões de Ga-

llardo, a maioria das subespécies

tem sua distribuição relacionada

com as grandes bacias hidrográfi-

cas embora subespécies diferentes

tenham sido encontradas em loca-

lidades próximas ou mesmo na

mesma localidade.

Nós, trabalhando independente-

mente, chegamos a conclusões se-

melhantes, o que vem confirmar o

acêrto do critério de Gallardo.

Entretanto, a obtenção de mate-

rial de regiões diversas daquelas

trabalhadas por Gallardo, princi-

palmente de zonas intermediárias,

veio mostrar que o problema da

distribuição geográfica das diver-

sas subespécies é por vêzes muito

complexo e de difícil solução dada

a exigüidade do material e dados

complementares.

Os esforços feitos nos últimos

anos no sentido de se obter séries

adequadas dêsse sapo nem sempre

foram compensados, pois se em al-

gumas ocasiões foram observados

exemplares em número espantoso,

em outras eram raros ou ausentes

dependendo usualmente da época

propícia, que é o início de grandes

chuvas.

Neste trabalho tratamos de ma-

terial da Amazônia, que embora

não muito variado e numeroso,

apresenta dados interessantes que

julgamos conveniente publicar.

Em 1960, recebemos do Dr. Ân-

gelo B. M. Machado, da Faculdade

de Medicina de Belo Horizonte, um

lote de exemplares por êle obtidos

na Cachoeira Uranduique, no rio

Mau, na região fronteiriça entre o

104

Atas do Simpósio sôbre a Biota Amazônica

Território Federal de Roraima e a

Guiana Inglesa. Êsse material de

pronto chamou a nossa atenção

por apresentarem os exemplares

uma cabeça particularmente larga,

diferenciando-se de todos os demais

que já tínhamos visto; o colorido

também, principalmente dos jo-

vens, era bastante característico.

Com o aparecimento do trabalho

de Gallardo, retomamos o estudo

do nosso material e verificamos que

os exemplares de Roraima são bas-

tante próximos à subespécie me-

rianae Gallardo, do rio Essequibo

na Guiana Inglêsa. O confronto de

nossos exemplares com parátipos

de merianae veio mostrar uma sé-

rie de diferenças constantes, pelo

que resolvemos descrevê-los como

subespécie nova.

Bufo granulosus nattereri subsp. n.

(Figs. 1-3, 7 e 8)

Holótipo — WCAB 7 . 484, macho

adulto, da Cachoeira Uranduique,

rio Mau, Território Federal de Ro-

raima, Brasil, colecionado em

VII — 1960 por Ângelo Machado.

Parátipos — WCAB 7.485-91,

machos adultos, colecionados jun-

tamente com o holótipo.

Diagnose — Uma subespécie de

Bufo gramdosus próxima a meria-

nae, caracterizada pela cabeça mui-

to curta e alta, paratóides muito

salientes, espaço interorbital bas-

tante largo e cristas cefálicas tu-

berculosas.

Descrição do holótipo — Cabeça

muito curta, expandida lateral-

mente. Focinho curto, saliente, um

pouco entumescido lateralmente.

Narinas ovaladas e largas, dispos-

tas obliquamente. As cristas cefáli-

cas são tôdas bem desenvolvidas,

de superfície tuberculosa. As pós-

-orbitais são mal perceptíveis; as

orbitais muito altas delimitando

um espaço interorbital que é mais

largo que a largura da pálpebra

superior; cantais e nasais bem de-

senvolvidas com as bordas tuber-

culosas; suborbitais salientes so-

bressaindo à margem lateral da ca-

beça, delimitando juntamente com

a crista maxilar um profundo sul-

co; pré e pós-orbitais bem desenvol-

vidas e salientes; cristas maxilares

bastante tuberculosas e salientes.

Tímpano quase vertical, de contor-

no pouco definido.

Paratóides curtas e muito sa-

lientes, de contorno nítido e des-

tacado e cobertas de espículas cô-

nicas.

Calosidades nupciais bem desen-

volvidas no polegar; calos carpais,

tarsais e pontas dos dedos e arte-

lhos enegrecidos.

O colorido dorsal apresenta-se

bastante contrastado. Sôbre um

fundo amarelado aparecem nume-

rosas manchas pretas, destacando-

-se um V invertido atrás da cabe-

ça e uma série de manchas for-

mando um M no sacro. O ventre

Volume 5

mostra manchas escuras apenas vi-

síveis lateralmente. Não há pro-

priamente uma linha vertebral

clara, porém as manchas escuras

estão interrompidas ao longo da

linha vertebral.

Medidas do holótipo — Compri-

mento total 55 mm; largura da ca-

beça 20; diâmetro ocular 6; lar-

gura da pálpebra superior 5; coto-

velo até a ponta do 3.° dedo 24;

(Zoologia) 105

fêmur 21; tíbia 21; calcanhar até

ponta do 4.° artelho 29.

Variação — Na série de paráti-

pos vê-se apenas variação quanto

ao colorido: alguns exemplares são

de fundo muito escuro e em razão

disso as manchas são pouco con-

trastadas. Em apenas um dos pa-

rátipos o ventre apresenta-se man-

chado esparsamente como no ho-

lótipo, pois nos demais as manchas

Figs. l a 3 — Cabeça do holótipo de Bufo granulosus nattereri subsp. n.;

figs. 4 a 6 — cabeça de Bufo granulosus merianae CAMNH ex series 46525-30).

cm

106

Atas do Simpósio sôbre a Biota Amazônica

são bem evidentes principalmente

na margem lateral. O comprimen-

to varia entre 50 e 53 mm com uma

média de 52 mm.

Juntamente com a série típica

recebemos 6 exemplares jovens

(WCAB 7493-98) cujo comprimen-

to varia entre 16 e 20 mm. Nestes

exemplares as cristas cefálicas ain-

da estão em parte incompletas po-

rém mostram-se bastante granulo-

sas. As cantais, as supraorbitais,

preorbitais e pós-orbitais são bem

evidentes; as maxilares e timpâni-

cas pouco visíveis e as infra-orbi-

tais ainda não formadas. O espaço

interorbital é completamente pla-

no e as paratóides bem delimitadas

e salientes, mesmo no exemplar

menor. O desenho dorsal também

é singular. Entre os olhos há uma

mancha triangular preta com o

vértice voltado para trás e unido a

ela uma mancha em forma de V in-

vertido. No sacro há uma grande

mancha preta dividida ao meio no

sentido longitudinal. Os flancos

apresentam-se quase pretos. A

gula é clara com vermiculação es-

cura e o peito e parte anterior da

abdômen quase preta com vermi-

culação amarelada.

Para comparação com a subes-

pécie merianae contamos com qua-

tro pará tipos da série AMNH

46.525-30 (3 machos e uma fê-

mea, adultos), (figs. 4-6).

Esta subespécie é dedicada a Jo-

hann Natterer, naturalista do Mu-

seu de Viena, que durante longos

anos colecionou na Amazônia no

comêço do século passado.

Fig. 7 — Vista dorsal do holótipo de

Bufo granulosus nattereri subsp. n.

(comprimento total 55 mm).

Bufo granulosus humboldti

Gallardo, 1965

Material examinado : Roraima,

3 exs. WCAB 18.782-84 — São

Marcos, rio Urariquera.

O encontro dessa subespécie co-

lombiana no Território de Roraima

é, até certo ponto, surpreendente,

ampliando de muito a distribuição

conhecida.

Os três exemplares concordam

com material que possuímos do Pa-

namá, inclusive no tamanho; são

machos adultos e medem 40, 41 e

44 mm.

Volume 5 (Zoologia)

107

\

108

Atas do Simpósio sôbre a Biota Amazônica

Em um dos exemplares (18.783)

a crista maxilar é bem menos de-

senvolvida que nos demais.

Bufo granulosus goeldii

Gallardo, 1965

Material examinado : Amapá,

4 exs. 2390-94 — Macapá, 1 ex.

2319 — Serra do Navio, 2 exs.

3094-95 — Pôrto Santana, 2 exs.

30953-54 — Mazagão. Pará, 4 exs.

30932-35 — Monte Alegre, 45 exs.

7356-400 — Maicuru. Amazonas,

4 exs. 30930-31, 30943, 30950

Manaus, 29 exs. 12039-67 — Mani-

coré. Rondônia, 25 exs. 10076-100

— Abunã. 74 exs. 11174-247 — Pôr-

to Velho.

O material do Amapá é o único

que apresenta diferenças em rela-

ção à forma típica que é da região

de Santarém. A cabeça é nitida-

mente mais curta e as cristas pós-

-orbitais mais evidentes, demons-

trando certos exemplares uma se-

melhança com merianae.

Bufo granulosus mirandaribeiroi

Gallardo, 1965

Material examinado-. Pará, 1 ex.

17946 — Alto rio Cururu, 5 exs.

40-42, 73, 8787 — Cachimbo, 3 exs.

16906-08 Conceição do Araguaia.

Maranhão, 1 ex. 7457 — Impera-

triz. Mato Grosso, 7 exs. 30936-42.

Barra do Tapirapé, 2 exs. 30944-45

— Xingu, 1 ex. 15164 — Rondonó-

polis, 4 exs. 15156-59 — Alto Gar-

ças.

É esta a subespécie que apresen-

ta até o momento os mais intrin-

cados problemas de distribuição.

Assim, os exemplares de Cachim-

bo e Alto rio Cururu são típicos re-

presentantes de mirandaribeiroi,

embora essas duas localidades per-

tençam à bacia do rio Tapajós. To-

dos os exemplares exceto um, apre-

sentam a linha vertebral clara bem

definida. A exceção é um jovem re-

cém-metamorfoseado de Cachimbo

com apenas 13 mm de comprimen-

to, cujo dorso é uniformemente es-

curo.

Os dois exemplares de Xingu

apresentam um problema análogo,

pois se assemelham em tudo a ma-

terial de Barra do Taripé e Con-

ceição do Araguaia, possuindo li-

nha vertebral clara bem definida.

Os 4 exemplares de Alto Garças

estendem a distribuição da subes-

pécie até às cabeceiras do Garças

que é um dos formadores do rio

Araguaia.

O exemplar de Rondonópolis

apresenta uma outra exceção pois

a região é da bacia do rio São Lou-

renço. O exemplar é semelhante

aos de Alto Garças inclusive quan-

to a detalhes de desenho.

O material acima citado, embo-

ra reduzido, sugere que nas cabe-

ceiras dos afluentes do Amazonas

Volume 5 (Zoologia)

109

que nascem no planalto central, a

distribuição das subespécies não

obedece mais à formação das ba-

cias hidrográficas, porém a outros

fatores como clima e principalmen-

te vegetação.

Tanto o Alto Cururu, como Ca-

chimbo e Xingu, são zonas de tran-

sição entre a floresta amazônica e

os campos cerrados do Brasil Cen-

tral, sendo sob êsse aspecto o Alto

Cururu mais amazônico que Ca-

chimbo e êste mais do que Xingu.

No caso do exemplar de Rondo-

nópolis observa-se caso análogo

pois embora a região pertença hi-

drogràficamente ao pantanal do

São Lourenço, fisiogràficamente é

muito semelhante a Alto Garças o

que explicaria a presença do mi-

r andar ibeir oi.

Expressamos aqui nossos agradeci-

mentos ao Dr. Richard Zweifel, do

American Museum of Natural History

de New York, e ao Dr. Antenor Leitão

de Carvalho, do Museu Nacional do Rio

de Janeiro, por sugestões e empréstimo

de material.

SUMÁRIO

O trabalho refere-se a distribui-

ção geográfica de Bufo granulosus

Spix, 1824, na Amazônia. Os estu-

dos do autor confirmam as conclu-

sões de Gallardo (1965) ao reco-

nhecer 14 subespécies. Além da des-

crição de uma subespécie nova, são

citados dados para B. g. humboldti,

B. g. goeldii e B. g. mirandaribeiroi.

Ilustrações e um mapa de distri-

buição de Bufo granulosus na Ama-

zônia brasileira são apresentados.

referência bibliográfica

gallardo, J. M., 1965, The species Bufo

granulosus Spix (Salientia, Bufo-

nidae) and its geographic varia-

tion. Buli. Mus. Comp. Zool, 134

(4) : 107-138.

Atas do Simpósio sôbre a Biota Amazônica

Vol. 5 (Zoologia): 111-125 — 1967

ESCORPIÕES, ARANHAS E ESCOLOPENDROMORFOS

DA AMAZÔNIA

WOLFGANG BÜCHERL

Instituto Butantan, São Paulo

A. ESCORPIÕES

O levantamento — que até esta

altura ainda deve ser considerado

provisório e insuficiente — das es-

pécies de escorpiões, encontradas

na biota amazônica, permite desde

já conclusões interessantes e eluci-

dativas.

Existem hoje no mundo inteiro

7 famílias, 22 subfamílias, cêrca de

95 gêneros e 710 espécies e na re-

gião neotropical, até aos altiplanos

que atravessam a linha leste-oeste

do México, 6 famílias. 25 gêneros

e cêrca de 210 espécies. Na biota

amazônica estão representadas, ao

que sabemos até hoje, 4 famílias,

9 gêneros e 28 espécies, sendo en-

dêmicas apenas:

15 espécies do gênero Tityus;

1 espécie do gênero Opistha-

canthus;

1 espécie (única) do gênero

Acanthothraustes;

4 espécies do gênero Proteo-

chactas e

1 espécie do gênero Broteas,

o que dá um total de 22 espécies

autóctones, pertencentes a apenas

5 gêneros e 3 famílias. O único gê-

nero realmente apenas amazônico

é Acanthothraustes, os outros qua-

tro mantêm sua área central de

dispersão fora da hylaea, tendo

apenas uma ou outra espécie con-

seguido tomar pé na Amazônia, de-

senvolvendo aí especiação própria.

1) Espécies endêmicas da biota

AMAZÔNICA

1>. Tityus amazonicus Giltay, . . .

1928 — capturado em Óbidos

e Belém;

2) T. cambridgei Pocock,1897 —

Amapá, Belém, Óbidos, Santa-

rém. Manaus, Teffé, Tabatin-

ga, Guianas e Venezuela (Sil-

vestre, em lugares elevados,

isentos das enchentes, o escor-

pião mais freqüente da re-

gião) ;

3) T. clathratus Koch, 1845 —

Pará, sem localização precisa;

112

Atas do Simpósio sobre a Biota Amazônica

4) T. discrepans (Karsch, 1879)

— Amapá ;

5) T. magnimanus Pocock, 1897

— Marajó;

6) T. metuendus Pocock, 1897 —

Pará (sem local de captura do

tipo) , Tabatinga;

7) T. paraensis Krpln. 1896 —

Pará (sem localidade) ;

8) T. silvestris Poc., 1897 — San-

tarém.

Espécies muito raras, mal descri-

tas ou conhecidas apenas por um

único exemplar:

9) Tityus asthenes Poc., 1893 (só

1 fêmea) — Baixo Amazonas;

10) T. bispinosus (Pessoa, 1934)

(só 1 macho) — Santarém;

11) T. carvalhoi M. L., 1945 (só 1

fêmea) — Tapirapés;

12) T. duckei Borelli, 1910 (só 1

macho) — Pará (sem locali-

dade) ;

13) T. evandroi M. L., 1945 (só 1

fêmea) — Piratuba;

14) T. sampaiocrulsi M. L., 1931

(só 1 macho) — Rio Cuminá;

15) T. strandi Werner, 1939 (só 1

fêmea) — Sacambu.

16) Opisthacanthus cayaporum

Vellard, 1932 — florestas do

Xingu;

17) Acanthothraustes brasiliensis

(M. L., 1931) (só 1 macho) —

Belém;

18) Proteochactas porosus Poc.,

1900 — Roraima;

19) P. granosus Poc., 1900 — Ro-

raima;

20) P. goujei Vellard, 1932 — (só

1 fêmea) — Xingu (Caiapós) ;

21) P. parvulus Poc., 1897 — San-

tarém, Belém;

22) Broteas paraensis Simon, 1880

— Belém e rio Xingu.

2. Espécies neotrópicas equato-

riais, QUE PENETRAM NA BIOTA

Amazônica

1) Rhopalurus laticauda Thorell,

1876 — centro de dispersão:

Nordeste brasileiro, Ceará;

com dispersão pelos altiplanos

do Amapá, das Guianas e Ve-

nezuela; Roraima;

2) Rhopalurus pintoi M. L., 1933

— rio Tacutu, Amazonas;

3) Diplocentrus g r u n dia chi

Karsch, 1880 — Venezuela,

Antilhas; 1 exemplar em

Belém;

4) Chactas vanbenedeni Gervais,

1844 — Colombia, Venezuela,

com penetração pela Hylaea

no nor-noroeste;

5) Teuthraustes amazonicus Si-

mon, 1880 — Ecuador, Peru

com penetração ao longo do

alto Solimões;

6) Broteas gervaisi Pocock, 1893

— Venezuela, Guianas, captu-

rado já às margens dos rios

Cuminá, Aurá e Tapirapés.

Volume 5 (Zoologia)

113

3. Discussão comparativa da es-

CORPIO-FAUNA MUNDIAL, NEOTRÓPICA

E AMAZÔNICA

382 espécies, pertencentes a cêr-

ca de 60 gêneros e 5 subfamílias,

formam a família mais importante

dos escorpiões, Buthiãae. E’ a úni-

ca entre tôdas que conseguiu fir-

mar e manter-se nos 5 continen-

tes. A subfamília Buthinae com-

preende cêrca de 54 gêneros e nada

menos de 230 espécies, tôdas elas

do Velho Mundo (África do Norte

desde as costas do Atlântico até o

mar Vermelho e a índia, África do

Sul, Ásia Menor, países do Medi-

terrâneo). A subfamília Ananteri-

nae com apenas 3 gêneros e 14 es-

pécies só, tem dispersão geográfi-

ca assaz curiosa: Ananteroides,

com 1 só espécie, é da Guiné por-

tuguesa, na África; Ananteris com

3 espécies é exclusivamente neotro-

pical, preferindo as espécies um

clima sêco, mesmo quando já com

temperaturas subtropicais, como

existe no Paraguai, no sul de Mato

Grosso, na Argentina ocidental,

nos vales pré-andinos do Ecua-

dor até a Venezuela; Isometrus, fi-

nalmente, abrange 11 espécies, tô-

das da região indo-australiana.

Apenas a espécie Isometrus ma-

culatus (De Geer, 1778) é tropical

cosmopolita (Austrália, índia, Áfri-

ca, América do Sul) , sendo presen-

te também em Punã, Belém, San-

tarém e Manaus. A possibilidade

de uma dispersão passiva existe.

As 40 espécies de Centruroides

(Centruroidinae) apresentam sua

maior densidade numérica nos alti-

planos secos do México, irradiando

daí para o norte (Sul dos EUA) e

para o sul (América Central, al-

gumas ilhas das Antilhas) sem

atingir propriamente dito o extre-

mo norte da zona neotropical. Seu

antagonista biológico é o gênero

Rhopalurus (Rhopalurinae) , neo-

tropical, das regiões secas do Nor-

deste brasileiro (epicentro de den-

sidade) com dispersão esporádica

pelos altiplanos das Guianas e da

Venezuela, porém quase sempre

nas imediações de portos maríti-

mos, com exceção de R. laticauda,

encontrado uma vez no sopé da

serra de Roraima. Onde termina a

dispersão de Centruroides, começa

a de Rhopalurus. De fato, são es-

corpiões com fácies morfológico

quase idêntico, sendo em certos ca-

sos extremamente difícil determi-

nar-se o gênero.

Tityus ( Tityinae , Buthiãae) fi-

nalmente é o gênero neotropical

mais abundante em espécies. Con-

quistou pràticamente tôda a Amé-

rica do Sul, com exceção do Chile.

O centro de densidade (T. serrula-

tus) é formado por certas regiões

de Minas Gerais (Nova Era, arre-

dores de Belo Horizonte, Monleva-

de), Goiás, Estado de São Paulo

(regiões de Ribeirão Prêto, Apare-

cida) e para T. bahiensis a vasta

área da margem direita do São

8 — 37 145

114

Atas do Simpósio sôbre a Biota Amazônica

Francisco até Mato Grosso e Para-

guai, ocorrendo o desvio este-oeste

justamente no Estado de São Pau-

lo em direção a Mato Grosso. Estas

duas espécies devem compreender

mais de 4/5, pois as demais 71 es-

pécies são relativamente raras,

com 15 espécies amazônicas. Colo-

rido e sub-regiões geográficas indi-

cam que as 73 espécies são reduzí-

veis a 4 subgêneros:

a) escorpiões silvestres equato-

riais, escuros; T. cambridgei da

hylaea amazônica pode servir como

protótipo;

b) escorpiões claros ou, quando

escuros, com pernas amareladas,

de uma côr só: T. serralatus e ba-

hiensis e afins;

c) escorpiões, cujo dorso é per-

corrido por faixas longitudinais:

grupo trivittatus;

d) escorpiões com corpo e pernas

irregularmente manchados.

A família Diplocentridae abran-

ge as subfamílias Nebinae com 2

gêneros e cada um apenas com 1

espécie da Síria, Arábia, Palestina

e Diplocentrinae. Esta, com 3 gêne-

ros que formam como que uma

ponte entre a zona sul da região

neártica e a zona norte da neotro-

pical. As duas espécies do gênero

Didynocentrns vão do México até

a América Central, a espécie úni-

ca de Oechis é da ilha Antigua nas

Antilhas e as 5 espécies de Diplo-

centrus vão desde as Antilhas até

a Venezuela, nos arredores dos por-

tos marítimos. Se não houve en-

gano na rotulação das peças do

Museu Britânico, houve um acha-

do de D. grundlachi em Belém.

148 espécies da África, Austrália,

de Madagascar, Etiópia, índia, Cei-

lão, Sumatra formam os 16 gêneros

e as 6 subfamílias de Scorpionidae.

Curioso é o gênero Opisthacan-

thus, cujo centro de dispersão com

17 espécies é a África. Entretanto,

2 espécies são neotropicais, elatus

das Antilhas, do Panamá e da Co-

lômbia e cayaporum relativamente

freqüente sob cascas de árvores,

sob troncos das florestas do Xingu,

na serra dos Caiapós.

A família dos Bothriuridae

abrange 9 gêneros e 55 espécies,

tôdas exclusivamente sul-america-

nas. O epicentro de freqüência está

na Argentina, no Uruguai e nos Es-

tados do Rio Grande do Sul e Pa-

raná, daí se processando a disper-

são em direção norte até o nordes-

te semi-árido do Brasil (Ceará) e

em direção aos andes, com gêneros

andinos que chegam até o Equa-

dor e a Colombia, sem que haja in-

vasão de hylaea amazônica.

A família dos Chaerilidae com-

preende apenas 2 gêneros com 12

espécies, tôdas da sub-região indo-

-malaia.

Em Vejovidae foram estabeleci-

das 6 subfamílias com 13 gêneros

e 54 espécies. As 19 espécies de Ve-

jovis e Hadrurus, Vejovinae, se su-

perpõem geogràficamente à disper-

Volume 5 (Zoologia)

115

são de Centruroides; os gêneros

Physoctonus e Caraboctouus, com

uma espécie cada um, são neotro-

picais, da Paraíba no nordeste do

Brasil e do sul do Chile respecti-

vamente; o gênero Hadruroides,

Caraboctoninae , contém 5 espécies

neotropicais, dos vales dos Andes

desde Chile até Equador, mas sem

penetração pelas florestas amazô-

nicas.

A família Chactidae contém as

subfamílias Euscorpioninae com 2

gêneros e 11 espécies da Europa,

em tôrno do Mediterrâneo, Mega-

corminae com 2 gêneros e 3 espé-

cies do México (Vera Cruz) e Chac-

tinae, com 7 gêneros exclusivamen-

te neo-tropicais, abrangendo 32 es-

pécies. O gênero Chactas contém

14 espécies andinas, desde o sul do

Peru até o oeste da Venezuela, pe-

netrando C. vanbenedeni pelo oeste

da Amazônia; as 13 espécies de

Teuthraustes acompanham a dis-

tribuição geográfica de Chactas,

embora mais freqüentes nos vales

andinos mais quentes e com abun-

dante vegetação, tendo sido encon-

trado T. amazonicus no oeste da

hylaea; as 7 espécies de Broteochac-

tas formam como que a continua-

ção dos primeiros dois gêneros,

ocorrendo desde Colômbia, Pana-

má, Venezuela, Trinidad, Guianas

até o Atlântico, tendo sido encon-

tradas 4 espécies na Amazônia

(Roraima, serra dos Caiapós, San-

tarém e Belém; o mesmo habitat

mantém o gênero Broteas, com 5

espécies amazônicas no sentido

amplo (Trinidad, Venezuela, Guia-

nas, ao longo dos rios Cuminá,

Aurá, Tapirapés, Xingu e cidade

de Belém).

B. ARANHAS

I. ARANHAS CARANGUEJEIRAS

Cêrca de 1615 espécies no mun-

do inteiro; 578 espécies na região

neotropical; 310 espécies na sub-

-região brasileira; 59 espécies na

biota amazônica.

1 . Actinopodidae — 3 gêneros —

2 exclusivamente neotropicais; 41

espécies.

1. Actinopus Perty, 1833 — 28

espécies exclusivamente neo-

tropicais, desde a Argentina

até o Panamá e Trinidad; 3 es-

pécies amazônicas: 1. A. nat-

tereri Auss., 1871) — Rio Ne-

gro — uma só fêmea, 12 mm;

2. A. piceus (Auss., 1871) —

Amazonas — uma só fêmea,

8,70 mm; 3. A wallacei. F.

Cambr., 1896 — Santarém —

uma só fêmea, 25 mm.

2. Missuleiia Walck., 1805 — 11

espécies — região zoo-geográ-

fica australiana.

3. Neocteniza Poc., 1895 — 2 es-

pécies neotropicais (Guatema-

la e Guiana ingl.) : 4. N. scla-

teri Poc., 1895 — Guiana ingl.

— uma só fêmea; 15 mm.

116

Atas do Simpósio sobre a Biota Amazônica

2 . Paratropididae — Com apenas

3 gêneros e 5 espécies exclusiva-

mente neotropicais, amazônicas:

4. Anisaspis Sim., 1891 — uma

só espécie: 5. A. tuberculata

Sim., 1891 — S. Vicente e Pará

— uma só fêmea, 6 mm .

5. Anisaspoides F. Cambr., 1896

— com uma só espécie: 6. A.

gigantea F. Cambr., 1896 —

Baixo Amazonas (Breves) —

uma só fêmea, 12,75 mm.

6. Paratropis Sim., 1889 — com

três espécies: 7. P. papilligera

F. Cambr., 1896 — Santarém

— macho e fêmea, 12,70 mm;

8. P. sanguínea M. L., 1923 —

Amazonas (rio Juruá) — uma

só fêmea, 14 mm; 9. P.

scruposa Sim., 1889 — Peru

(Pebas) — uma só fêmea,...

10 mm.

3. Ctenizidae — com 99 gêneros

e 546 espécies:

Ctenizinae — com 91 gêneros e

512 espécies das regiões zoo-geo-

gráficas paleártica, etiópica, afri-

cana, austral, oriental, neártica

e neotropical. 50 espécies neotro-

picais; 27 espécies brasileiras;

0 espécies amazônicas.

Idiopinae — com 2 gêneros e 18

espécies exclusivamente neotro-

picais e sul-americanas:

7. Idiops Perty, 1833 — 12 espé-

cies: 10. I. petitii (Guérin,

1838) — Santarém — um ma-

cho só, 30 mm; 11. I. santa-

remia (F. Cambr., 1896) —

Santarém — uma fêmea só,

16 mm.

8. Pseudidiops Sim., 1889 — 6 es-

pécies: 12. P. opifex Sim., 1889

— Guiana fr. (Caiena) — uma

fêmea só; 17 mm.

Bhachybothriinae — com 6 gêneros

e 16 espécies, sendo 5 gêneros e

15 espécies da região neártica e

1 gênero com uma espécie do

Japão.

4. Migidae — com 11 gêneros

e 45 espécies das regiões austral,

sul-africana, Madagascar, sendo

neotropical apenas o gênero Cala -

thotarsus com a espécie única co-

ronatus Sim., 1903 (Chile), uma só

fêmea.

5. Dipluridae — Com 50 gê-

neros e 210 espécies das regiões

sul-africana, australiana, neotropi-

cal, oriental, paleártica, neártica,

tendo 22 gêneros representantes

neotropicais; 79 espécies neotropi-

cais sendo 30 brasileiras. Espécies

amazônicas:

9. Diplura C. L. Koch, 1850 —

18 esp. neotropicais: 13. D.

fallax M. L., 1926 — rio Juruá

— uma fêmea só.

10. Eudiplura Sim., 1892 — com a

única espécie: 14. E. rogenho-

feri (Auss., 1871) — Amazo-

nas (norte) e Venezuela.

Volume 5 (Zoologia)

117

11. Fufius Sim., 1888 — com 7 es-

pécies, sendo 1 de Guatemala,

1 do Equador, 2 da Bolívia e

as restantes amazônicas: 15.

F. albovittatus (Sim., 1891) —

Manaus — um macho só; 8

mm; 16. F. auricomis (Sim.,

1891) — Santarém — um

macho só; 12,50 mm; 17. F.

funebris Vellard, 1924 — Ara-

guaia.

12. Harmonicoii F. Cambr., 1896

— 1 espécie do Equador, 1 do

Rio de Janeiro e 18. H. rufes-

cens F. Cambr., 1896 — San-

tarém — uma fêmea só; 27

mm.

13. Thalerothele Bertk., 1880 —

com 6 espécies de São Paulo,

Rio, Minas, sendo amazôni-

cas: 19. Th. fasciata Bertk.,

1880 — Belém — só uma fê-

mea; 14 mm; 20. Th. nigra F.

Cambr., 1896 — Santarém; só

uma fêmea só; 30 mm.

sanguínea (F. Cambr.) — San-

tarém; só uma fêmea; 24 mm.

uma fêmea só; 30 mm.

14. Trechona C. L. Koch, 1850 —

com 5 espécies exclusivamente

sul-americanas, sendo uma do

Rio de Janeiro, outra de São

Paulo, uma terceira de Bogo-

tá e as seguintes duas com

área de dispersão pelas Guia-

nas até Amazonas, Pará até

São Paulo: 22. T. venosa

(Latr., 1830), 30,00 mm; 23.

T. lycosiformis (C. L. Koch) —

uma fêmea só; 30 mm.

15. Ischnothele Auss., 1875 —

com 18 espécies da África, ín-

dia, do México, Guatemala,

Nicarágua, Costa Rica, Pana-

má, Cuba, Venezuela: 24. I.

guyanensis (Walck., 1837) —

de Costa Rica, Panamá, Guia-

nas até Amazonas; 25. I. sie-

mensi F. Cambr., 1896 —

autóctone de Pará — uma só

fêmea; 18 mm.

6. Atypidae — com 2 gêneros e

18 espécies das regiões paleárticas,

indo-austral, neártica e africana.

Ausente na neotropical.

7. Barychelidae — com 58 gê-

neros no mundo inteiro, com 150

espécies; 26 gêneros neotropicais,

com 55 espécies; 14 gên. brasilei-

ros, com 23 espécies; 6 gên. ama-

zônicos, com 8 espécies:

16. Cyrtogrammomma Poc., 1895

— com uma só espécie: 26.

C. monticola Poc., 1895 — Ro-

raima.

17. Homoeoplacis Sim., 1892, com

duas espécies só: 27. H. austeni

F. Cambr., 1896: Manaus —

um macho só; 11 mm; 28. H.

pentodon Sim., 1892: Baixo

Amazonas (S. Paulo?).

18. ldiophthalma Cambr., 1877,

com seis espécies, sendo 2 de

Quito, 1 da Colombia, 1 de San

Esteban: 29. 1. pantherina

Sim., 1889: San Estevan> Va-

118

Atas do Simpósio sobre a Biota Amazônica

lenda : Venezuela e no alto rio

Negro: Amazonas; 30. I ama-

zônica Sim., 1889: Teffé: Ama-

zonas ocidental — uma fêmea

só; 11 mm.

19. Strophaeus Auss., 1875, com

uma só espécie: 31. St. kochii

(Cambr., 1870) — Amazonas:

Pebas (Peru) : um macho só;

14 mm.

20. Trichopelma Sim., 1888, com

três espécies, sendo uma da

Bahia, a segunda da ilha de

S. Domingos, e: 32. T. illeta-

bile Sim., 1888 — Teffé; uma

fêmea só; 20 mm.

21. Diplothelopsis Tullgren, 1905,

com cinco espécies, sendo 4 da

Argentina: 33. D. hastatus

M. L., 1923 — Alto Juruá; uma

fêmea só; 18 mm.

8. Pycnothelidae — com 3 gê-

neros exclusivamente sul-america-

nos, com apenas um total de 4 es-

pécies do oeste da Argentina (Cór-

doba, Catamarca, Colonia Dora) e

Pycnothele perditus Chamb., 1917,

espécie genotípica, do Rio de Ja-

neiro.

9. Theraphosidae — com 7 sub-

famílias:

Ischnocolinae — com 45 gêne-

ros nas regiões neártica, africa-

na, neotrópica, paleártica, indo-

austral, com 154 espécies; com

33 gêneros e 101 espécies neotro-

picais; com 27 gêneros e 31 es-

pécies brasileiras, sendo da Ama-

zônia apenas as seguintes espé-

cies:

22. Cyriocosmus Sim., 1903, com

1 espécie de Trinidad e 3 da

hylaea: 34. C. elegans (Sim.,

1889) — Vale do Orinoco; 35.

C. nigriventris M. L., 1939 —

Falcon: Venezuela; 36. C. sel-

latus (Sim., 1889) — Amazo-

nas — Fonteboa, S. Paulo de

Olivença — 14 mm.

Grammostolinae — com 18 gê-

neros, exclusivamente neotro-

picais e neárticos; 90 sp. com 10

gêneros exclusivamente sul-ame-

ricanos, 2 intermediários; com

74 espécies neotropicais, sendo

52 sul-americanas, das quais 20

brasileiras, dos Estados do sul do

Brasil.

Theraphosinae — com 11 gê-

neros sul-americanos; apenas

Phormictopus tem representan-

tes nas Antilhas até a Califór-

nia; com 112 espécies:

23. Acanthoscurria Auss. 1871,

com 37 espécies no Brasil (Est.

S. Paulo, Mato Grosso, Ceará,

Minas Gerais, Paraíba, Rio

Grande do Norte, Rio Grande

do Sul, Maranhão), Uruguai,

Argentina, Bolívia, 1 espécie

nas Antilhas: 37. A. brockle-

hursti F. Cambr., 1896: Pará;

uma fêmea só; 60 mm; 38.

A. jerina Sim., 1892. Tefé; um

macho só; 40 mm; 39. A.

jracta Chamb., 1917; Pará; um

Volume 5 (Zoologia)

119

macho só; 50 mm; 40. A. geni-

culata (C. L. Koch, 1842) , San-

tarém e Rio Branco; 70 mm;

41. A. minor Auss., 1871;

Guiana; um macho só; 42. A.

tarda Poc., 1903; Tefé; uma

fêmea só; 62 mm; 43. A.

theraphosoides (Auss., 1871) ;

Maranhão; um macho só; 40

mm; 44. A. xinguensis T.

Costa, 1960, Alto Xingu; 50

mm.

24. Lasiodora C. L. Koch, 1850 —

21 espécies de São Paulo, Rio

de Janeiro, Minas Gerais, Es-

pírito Santo, Bahia, Paraíba,

Sta. Catarina, sendo duvidosa

a ocorrência em Montevidéu e

na Argentina: 45. L. Lakoi

M. L., 1943 — rio Purus; um

macho só; 40 mm.

25. Megaphobema Poc., 1901 —

uma espécie: 46. M. robusta

(Auss., 1875) — Colômbia e

Uaupés, no rio Negro.

26. Sericopelma Auss., 1875 — 3

espécies, sendo duas do Pana-

má: 47. S. fallax M. L., 1923

— rio Juruá; um macho só;

58 mm.

27. Theraphosa Thorell, 1870 —

uma espécie: 48. Th. blondii

(Latreille, 1804) — Venezuela

oriental, Guianas, Amapá

(Serra do Navio) — uma fê-

mea só; 80 mm.

28. Trasiphoberus Sim., 1903 —

uma só espécie: 49. T. parvi-

tarsis Sim., 1903 — Teffé;

52 mm.

29. Xenesthis Sim., 1891 — com

3 espécies — Colômbia até

Venezuela: 50. X. immanis

(Auss., 1875) — Colômbia Oci-

dental, Alto Rio Negro; 75 mm.

51. X. monstruosa Poc., 1903

— o mesmo habitat; uma fê-

mea só.

A viculariinae — Com 7 gêne-

ros, sendo 3 africanos e os res-

tantes neotropicais; com 11 es-

pécies africanas e 47 neotropi-

cais, sendo da hylaea amazônica

as seguintes:

30. Avicularia Lamarck, 1818 —

com 32 espécies: Amazônia,

Guianas, Trinidad, Pôrto Rico,

Califórnia, Panamá, Colombia,

Paraíba, Maranhão: 52. A. an-

cylochyra M. L., 1923 — rio

Tapajós — uma fêmea só; . . .

35 mm; 53. A. a. avicularia

(Linn., 1758) — Guianas,

Trinidad, tôda a Amazônia;

43 mm; 54. A. bicegoi M.

L. , 1923 — Manaus; uma

fêmea só; 36 mm; 55. A.

cuminami M. L., 1930 — rio

Cuminá; sem descrição da es-

pécie; 56. A. rufa Vellard,

Schiap. & Gerschm., 1945 —

Alto Gy — Pará; uma fêmea

só; 37 mm; 57. A. juruensis

M. L., 1923 — rio Juruá; uma

fêmea só; 50 mm; 58. A.

120

Atas do Simpósio sôbre a Biota Amazônica

metallica (Auss., 1875) —

Guiana hol.

Selenocosmiinae — com 16 gêneros

e 127 espécies, principalmente

nas regiões indo-austral, e afri-

cana, sendo sul-americanos ape-

nas os 2 gêneros Ancylochiros

M. L., 1920 com uma espécie de

Minas Gerais, e Ephebopus:

31. Ephebopus Sim., 1892 — com

3 espécies do Equador e Bra-

sil: 59. E. murinus (Walck.,

1873) — Santarém — 1 fê-

mea só; 55 mm.

Eumenophoriinae — com 18 gêne-

ros com 56 espécies exclusiva-

mente africanas, sendo apenas

Erythropoicila plana da Vene-

zuela — um macho só.

Resumindo a ocorrência de Ara-

nhas Caranguejeiras na hylaea

amazônica, temos a constatar

ainda, a título de ensaio pré-

vio, os seguintes fatos interessan-

tes sob o ponto-de-vista zoogeo-

gráfico:

As representantes da família

Paratropididae só foram encontra-

dos na biota amazônica, enquanto

que as famílias Migidae, Atypidae

e Pycnothelidae não têm nenhum

habitante amazônico, embora os

Picnotelídeos fôssem exclusivamen-

te sul-americanos.

Dos numerosos Ctenizíneos com

27 espécies brasileiras, nenhuma

foi assinalada para a Amazônia,

o mesmo valendo no tocante aos

Ischnocolíneos e Gramostoloíneos,

com apenas 3 espécies amazônicas

entre os primeiros e nenhuma nos

últimos; entre os Terfosíneos há

um espécime só de Lasiodora, cêr-

ca de dez exemplares apenas

de Acanthoscurria. Megaphobema,

Theraphosa, e Xenesthis são, en-

tretanto, exclusivamente amazô-

nicos, embora sempre raríssimos,

constituindo as caranguejeiras de

maiores proporções de nossa terra.

O gênero Avicularia deve ser con-

siderado como essencialmente ama-

zônico, podendo haver, em certos

locais, dezenas de exemplares, que

vivem geralmente trepados sôbre

arbustos e mesmo árvores (palmei-

ras), nidificando aí mesmo.

Como testemunho da precarie-

dade das colheitas até agora fei-

tas na imensa hylaea ressaltam os

poucos locais de captura e ainda

o fato, de que das 59 espécies au-

tóctones da Amazônia nada menos

de 41 espécies estão conhecidas

apenas de um único exemplar (o

tipo) , nada se sabendo sôbre o sexo

oposto.

II . Aranhas verdadeiras

A limitação dêste trabalho não

permite a inclusão das aranhas

verdadeiras da hylaea, pois já agora

são certamente mais de 1.000 es-

pécies, distribuídas sôbre várias

centenas de gêneros e dezenas de

famílias. Além do mais, os nossos

Volume 5 (Zoologia)

121

conhecimentos são toldados pelos

poucos locais de captura, arrolados

até agora.

C. ESCOLOPENDROMORFOS

Com duas famílias.

1 . Scolopendridae — com duas

subfamílias.

Scolopendrinae — com duas

tribus.

a) Scolopendrini — com 7 gêne-

ros e cêrca de 163 espécies.

1. Scolopendra L., 1758 — com

cêrca de 60 espécies em todos

os países tropicais e de clima

subtropical; 18 espécies neo-

tropicais. 1. S. morsitans L.,

1758; tropical cosmopolita: S.

morsitans amazônica Bücherl,

1946 — Manaus, 86 mm;

2 . S. alternans Leach, 1805 —

Venezuela, Orinoco, Manaus

190 mm; 3. S. arthrorhab-

doides Ribaut, 1914 — Co-

lômbia, Amazonas: Naupés

70 mm; 4. S. gigantea gigan-

tea L., 1758 — América Cen-

tral, Venezuela, Amazonas

(Rio Branco e Rio Negro) 270

mm; 5. S. angulata angulata

Newp., 1844 — o mesmo habi-

tat de gigantea 170 mm; S.

angulata moojeni Bücherl,

1941 — rio Tapirapés 170 mm;

6. S. v. viridicornis Newp.,

1844 — Argentina, Uruguai,

Paraguai, Bolívia, Guianas,

Venezuela, Amazonas, Pará e

todo o resto do Brasil 200 mm;

7. S. v. viridis Say, 1821 —

desde o México até a Amazô-

nia 130 mm; S. v. polymor-

pha Wood, 1861 — o mesmo

habitat; 130 mm.

2. Trachycormocephalus Krpln.,

1903 — com 8 espécies da Áfri-

ca até a índia; ausente na re-

gião neotropical.

3. Arthrorhabdus Poc., 1891 —

com 4 espécies, sendo 1 da Aus-

trália, 1 da África do Sul,

1 do México e: 8. A. spinifer

(Krpln., 1903) de Belém do

Pará; 35 mm.

4. Cormocephalus Newp., 1844 —

com cêrca de 92 espécies tropi-

cal - cosmopolitas (Nova Ze-

lândia, Austrália, Madagascar,

África, Ceilão, Filipinas, índia,

Mediterrâneo, América do Nor-

te, Central e Sul) : 9 . C. me-

diosulcatus Att., 1928: ilha de

Marajó; 52 mm; 10. C. un-

gulatus (Mein., 1886) : Haiti,

Venezuela, Manaus, Recife;

45 mm; 11. C. brasiliensis

Humb. — Sauss., 1870 — Ve-

nezuela, Manaus; 24 mm;

12. C. impressus neglectus

(Chamb. 1914) — rio Madeira;

60 mm; 13. C. amazonae

(Chamb., 1914): Manaus

43 mm.

5. Campylostigmus Rib., 1923 —

com 5 espécies da Nova Cale-

dónia.

122

Atas do Simpósio sôibre a Biota Amazônica

6 . Rhoda Mein., 1886 — com 2 es-

pécies brasileiras (Bahia, Per-

nambuco, Santarém): 14. Rh.

thayeri Mein., 1886: Santarém

75 mm.

7. Scolopendropsis Brandt, 1841

— com 1 espécie de Salvador,

Bahia.

b) Asanadini — com 2 gêneros e

6 espécies:

8. Asanada Mein., 1886 — com 4

espécies: índia, Nova Guiné,

Filipinas, Palestina, África.

9. Pseudocryptops Poc., 1891 —

com 2 espécies: índia, Abissí-

nia.

Otostigminae — com 2 tribus:

a) Otostigmini — com 6 genê-

ros; todos os países tropicais e

subtropicais .

10. Otostigmus Por., 1876 — com

72 espécies: índia, Ceilão, No-

va Guiné, Bornéo, Burma, Ce-

lebes, Java, Madras, Sumatra,

Austrália, África, China, Co-

lômbia, Argentina, Uruguai,

Brasil, América Central, Méxi-

co, sendo da Amazônia as se-

guintes: 15. O. pococki Krpln.,

1903 — Margem esquerda do

Baixo Amazonas, Amapá

90 mm; 16. O. rex Chamb.,

1914 — Santarém (Taperi-

nha); Madeira - Mamoré

90 mm; 17. O. inermis Por.,

1876 — Cordilheira central e

oriental da Colômbia, Vene-

zuela, Buenos Aires — Rio Ne-

gro, Amazonas 54 mm; 18.

O. amazonae Chamb., 1914 —

Manaus 45 mm; 19. O. ca-

sus Chamb., 1914 — Madeira-

Mamoré 57 mm; 20. O. la-

tipes Bücherl, 1954 — Rio das

Mortes — Araguaia 31 mm;

21. O. tidius Chamb., 1914 —

Manaus 14,50 mm; 22. O. goel-

dii Brõl., 1898 — Belém, Vene-

zuela 24 mm; 23. O. clavi-

fer Chamb., 1921 — Guiana

Inglesa, Amazonas 60 mm;

24. O. brunneus Chamb., 1921

— Guiana Inglesa 32 mm;

25. O. suitus Chamb., 1914 —

Madeira — Mamoré 55 mm.

11. Digitipes Att., 1930 — com 1

espécie do Congo.

12. Alipes Imhoff, 1854 — com 5

espécies da África.

13. Ethmostigmus Poc., 1898 —

com 15 espécies: Etiópia, ín-

dia, Austrália.

14. Rhysida Wood, 1862 — com

18 espécies das regiões indo-

-australiana, etiópica e neo-

tropical e neártica, tendo re-

presentantes na Amazônia as

seguintes: 26. R. ceie ris

(Humb.-Sauss., 1876) — des-

de o México até a Argentina

— Manaus, Belém 70 mm;

27. R. nuda immarginata

(Por., 1876: índia, Filipinas,

Venezuela, Rio Negro 65 mm;

28. R. longipes longipes

'Volume 5 (Zoologia)

123

(Newp., 1845) : cosmotropical

— Manaus, Belém 90 mm.

15. Alluropus Silv., 1912 — com

1 espécie de Tonkin.

b) Arrhabdotini — com 1 gênero:

16. Arrhábãotus Att., 1930 — com

1 espécie de Bornéo.

2 . Cryptopidae — com 3 sub-

famílias:

Cryptopinae — com 4 gêneros

e 81 espécies:

17. Cryptops Leach., 1815 — com

70 espécies da Europa, Nova

Zelândia, Java, Burma, Nova

Guiné, África, Austrália, Su-

matra, índia, China. América

do Sul: Patagônia, Chile, Ar-

gentina, Bolívia, Brasil (Pa-

raíba do Norte) , América

Central (Cuba, Honduras,

Haiti), sendo da Amazônia as

seguintes: 29. C. furciferens

Chamb., 1921 — Guiana Ingle-

sa, ao longo do rio Demerara;

Brasil: Território do Rio Bran-

co; 15 mm.

18. Paracryptops Poc., 1891 —

com 4 espécies da índia, Nova

Guiné, Java, sendo 30. P. inex-

pectus Chamb., 1914., da Guia-

na inglesa; 16 mm.

19 . Anethops Chamb., 1902 — com

1 espécie da Califórnia.

20. Mimops Krpln., 1903 — com

2 espécies, sendo uma da Chi-

na, a segunda do Rio de Ja-

neiro.

Theatopsinae — com 2 gêne-

ros:

21. Theatops Newp., 1844 — com

3 espécies do sul dos Estados

Unidos, México e Mediterrâ-

neo.

22. Plutonium Cavanna, 1881 —

com 1 espécie da Sicília.

Dinocryptopiuae — com 6 gê-

neros e cêrca de 50 espécies.

23. Dinocryptops Crabill, 1958 —

com 5 espécies dos Estados do

Sul dos EUA, México. América

Central, Hylaea, América do

Sul e China. 30. D. miersii

(Newp., 1845) — desde os Esta-

dos Unidos até a Argentina,

Manaus, Belém, Óbidos.

24. Otocryptops Haase, 1887 —

com 8 espécies com a mesma

distribuição da espécie ante-

rior; 31. O. /. íerrugineus (L.,

1767) : África e as três Améri-

cas: Manaus, Belém; 32. O.

melanostomus (Newp., 1845) :

Filipinas, Nova Guiné, as três

Américas, Manaus, Belém,

Teffé.

25. Kethops Chamb., 1912 — com

1 espécie de Utah. E.U.A.

26. Kartops Archey, 1923 — com

1 espécie da Guiana Inglesa.

27. Tidops Chamb., 1915 — com

2 espécies, sendo uma de Gre-

nada, a segunda de Dunoon,

na Guiana Inglesa.

124

Atas do Simpósio sobre a Biota Amazônica

28. Newportia Gerv., 1847 — com

cêrca de 30 espécies, tôdas da

zona neotropica, sul do Méxi-

co, América Central, Antilhas,

norte da América do Sul, hy-

laea: 33. N. longitarsis sylvae

Chamb., 1914 — Madeira a-

-Mamoré; 34. N. bicegoi Brõl.,

1904 — Manaus; 35. N. ernsti

Poc., 1891 — Rio Negro; 36.

N. paraensis Chamb., 1914 —

Belém; 37. N. amazônica

Brol., 1904 — Manaus.

Entre as 420 espécies de Escolo-

pendromorfos estão representados

na biota amazônica 37 espécies,

principalmente dos gêneros Scolo-

pendra, Cormocephalus, Otostig-

mus, Rhysida e principalmente

Newportia.

RESUMO

Vinte e duas espécies endêmicas

da ordem dos Escorpiões, perten-

centes a 5 gêneros e 3 famílias, fo-

ram encontradas até agora na bio-

ta amazônica, a maioria delas ain-

da mal determinada, sendo os mais

comuns o Tityus cambridgei, o

Opisthacanthus cayaporum e o

Teuthraustes amazônicas. Os úl-

timos dois apresentam hábitos den-

drícolas, e o primeiro ocorre sôbre

elevações isentas das águas de

inundação.

Entre as 1615 espécies de Ara-

nhas Caranguejeiras, arroladas

para o mundo inteiro, figura a fau-

na amazônica com 59 espécies en-

dêmicas, pertencentes aos gêneros

Actinopus, Anisaspis> Anisaspoides,

Paratropis, Idiops, Fufius, Thalero-

thele, Idiophthalma, Acanthoscur-

ria, Megaphobema, Theraphosa,

Xenesthis e principalmente diver-

sas espécies de Avicularia. A maio-

ria, principalmente as que cons-

tróem tubos subterrâneos, munidos

de tampa do tipo de alçapão, vive

em locais isentos das inundações

periódicas, enquanto que as Avi-

cularias vivem sôbre arbustos e ár-

vores. Theraphosa blondii é a maior

aranha caranguejeira do mundo,

seguida pelos representantes de

Megaphobema e Xenesthis.

A fauna dos Escolopendromorfos,

com cêrca de 420 espécies no mun-

do inteiro, apresenta 37 espécies da

biota amazônica, pertencendo prin-

cipalmente aos gêneros Scolopen-

dra, Cormocephalus, Otostigmus e

principalmente Nexcportia. S. gi-

gantea com 27 cm de compri-

mento é o maior escolopendrídeo

do mundo.

SUMMARY

22 endemic species of Scorpions,

belonging to 5 genera and 3 fami-

lies are represented in the amazo-

na*1 biotope; the most common

are Tityus cambridgei, Opisthacan-

thus cayaporum and Teuthraustes

amazonicus.

The “tarantula” — like Mygalo-

morphe Spider fauna, with 1615

Volume 5 (Zoologia)

125

species in the whole world is pre-

sent in the hylaea with 59 ende-

mic species of the genera Actino-

pus, Anisaspis, Anisaspoides and

Paratropis, Idicps, Fufius, Thalero-

thele, Idiophthalma, Acanthoscur-

ria, Megaphobema, Theraphosa,

Xenesthis and chiefly Avicularia.

The aviculariid — Tarantulae are

dendrophyle during the most part

of their life; Theraphosa blondii

may be considered the biggest “Ta-

rantula’4 in the world, followed by

Xenesthis and Megaphobema.

The Scolopendromorphe — fau-

na with 420 species in the world

shows 37 species in the amazonian

biotope, distributed into the gene-

ra Scolopendra, Otostigmus, Cor-

mocephalus and chiefly Neivportia.

S. gigantea with 27 cm of body

length is the biggest Scolopendro-

morphe in the world.

BIBLIOGRAFIA

Attems, G„ 1930, Das Tierreich 54. Lief.

2. Scolopendromorpha, W. de Gruy-

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Borelli, a., 1910, Boll. Mus. Anat.

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Bücherl, W., 1941, Mem. Inst. Butantan

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Bücherl, W., 1943, Mem. Inst. Butantan

17: 19-28.

Bücherl, w„ 1946, Mem. Inst. Butantan

19: 135-158.

Bücherl, w., 1954. Mem. Inst. Butantan

26: 1-6.

Bücherl, W., 1959, Beitr. z. neotrop.

Fauna 1 (3) : 229-242.

Chamberlin, R. V., 1955, Lunds Univ.

Arskr., N. F. 51 (5) : 1-61.

De Geer, H., 1778, Mém. Inst. Aptères

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Gervais, E., 1844, Arch. Mus. Hist. Nat.,

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Giltay, L., 1928, Ann. Buli. soe. ent.

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Kraus, O., 1957, Senck. biol. 38 (5/6) :

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Atas do Simpósio sobre a Biota Amazônica

Vol. 5 (Zoologia): 127-139 — 1967

SIMULIIDAE DA AMAZÔNIA. III:

SOBRE O GÊNERO “SIMULIUM” LATREILLE, 1802

(Diptera, Nematocera) *

N. L. CERQUEIRA

Instituto Nacional de Pesquisas da Amazônia, Manaus

O papel que as espécies do gêne-

ro Simulium, há muito vêm de-

sempenhando na disseminação de

agentes etiológicos de doenças, tem

sido objeto de estudos em muitos

países. Entre nós, e particularmen-

te na Amazônia só recentemente

Cerqueira (1959) incriminou o Si-

mulium amazonicus Goeldi, 1905

como transmissor de microfilária

da Mansonella ozzardi, motivando

com isto, estudos mais acurados so-

bre esta e demais espécies do gê-

nero na região.

A tais estudos deveu-se não só

uma melhor definição da espécie de

Goeldi, já que as descrições exis-

tentes ofereciam dúvidas quanto às

características apresentadas pelo

autor, assim como também, por não

ser conhecido o seu procedimento

em relação à doença, nesta e nas

demais áreas da região.

Paralelamente, êsses estudos fo-

ram extendidos às espécies já des-

critas e às que foram sendo obti-

das no decorrer das pesquisas de

campo realizadas por nós desde. . .

1957.

Assim é que, até o momento, 10

diferentes espécies já foram encon-

tradas na região além do S. ama-

zonicum : o S. simplicicolor Lutz,

1910, proveniente de Rondônia

(Madeira-Mamoré, localidade ti-

po) cujo estudo taxinômico está

em pauta, eoS. subclavibranchium

Lutz, 1910 da qual não temos ain-

da material. Das demais, duas são

novas: S. goeldii, de material

oriundo de localidades do municí-

pio de Manaus e a outra S. fulvi-

notum de igual procedência, am-

bas descritas a seguir.

• Trabalho do Instituto Nacional de

Pesquisas da Amazônia, do Conselho

Nacional de Pesquisas (Diretor: Dr.

D j alma da Cunha Batista) , Divisão de

Pesquisas Biológicas (Diretor: Dr.

Mário Moraes) .

128

Atas do Simpósio sóbre a Biota Amazônica

HISTÓRICO

A família Simuliidae é represen-

tada na Amazônia por um único

gênero, Simulium Latreille, 1802.

Até 1905 nenhuma referência ha-

via sôbre as espécies dêsse gênero

no Brasil. Goeldi, nesse ano, dese-

jando resolver a questão da pre-

sença das mosquinhas abundantes

e incômodas existentes em algumas

áreas da região, resolveu a despei-

to da falta de um especialista, dar

algumas características a fim de

colocá-los no gênero Simulium e,

na mesma ocasião, descreveu o Si-

mulium amazonicum baseando-se

em exemplares de Tefé e alto Pu-

rus, no Estado do Amazonas.

Anteriormente, entrando em

contato com Lutz, a quem havia

enviado material para estudo, re-

cebia confirmação de que os exem-

plares eram de fato Simulium e

afirmava Lutz ter encontrado na

parte meridional do Brasil, mais

3 espécies dêste gênero além da-

quela; todavia nada publicara a

respeito do gênero. Dessa data em

diante nenhuma referência foi fei-

ta ao gênero na Amazônia até . . .

1910, quando Lutz, voltando a se

ocupar das espécies meridionais,

referiu-se a duas espécies encon-

tradas na Estrada de Ferro Madei-

ra-Mamoré, no Território de Ron-

dônia, das quais deu descrição

incompleta, a do S. simplicicolor,

com fêmea, macho e pupa e a do

S. subclavibranchium somente das

brânquias pupais.

A partir daquela data várias re-

ferências foram feitas ao S. ama-

zonicum, porém nenhum estudo foi

realizado com material da região,

até 1964, quando Cerqueira & Nu-

nes de Mello se ocuparam da es-

pécie de Goeldi descrevendo-a de

forma completa e apresentando

circunstanciado histórico pelo qual

fixaram-na, baseados em material

da localidade tipo (Tefé), ofere-

cendo ainda dados bionômicos, os

quais até então eram desconheci-

dos.

IMPORTÂNCIA

Os simulídeos são importantes

transmissores de doenças aos ani-

mais e ao homem. Vários são os re-

gistros existentes sôbre as diversas

zoonoses observadas na Europa e

América do Norte por êles veicula-

das.

O primeiro registro relacionado

com a transmissão de doenças ao

homem por êsses insetos, foi feito

por Blacklock em 1926 em que

dava o resultado de suas experiên-

cias sôbre a infestação por micro-

filária de Onchocerca volvulus

(Leuckart, 1893) pelo Simulium

damnosum Theob., 1903 em Sierra

Leone na África. Desta data em

diante vários são os autores que

em suas pesquisas têm se ocupado

do assunto.

Volume 5 (Zoologia)

129

Na América do Norte, Central e

do Sul certas espécies de Simulium

são incriminadas na transmissão

daquela doença africana.

Lane, em 1949, aproveitando um

trabalho de levantamento levado a

efeito em 1943, associou a presen-

ça do S. pruincsum com a ocorrên-

cia de casos de Penjigo foliaceo

(fogo selvagem) , em diversas loca-

lidades no Estado de São Paulo.

Recentemente, Anderson e cols.

isolaram virus tipo Leste de S. me-

ridionale, em Wisconsin, na Amé-

rica do Norte.

No Brasil, embora o gênero seja

conhecido desde o comêço do sé-

culo, os estudos têm sido dirigidos

mais para taxinomia das espécies

e não em pesquisas áo seu proce-

dimento, e só agora é que foi ve-

rificada na Amazônia o com-

prometimento de uma delas na

veiculação de microfilária da Man -

sonella czzardi. Contudo, dada a

distribuição das espécies em nosso

país é de se esperar que pesquisas

sejam feitas no sentido de conhecer

se outros simulideos possam estar

incriminados nessa ou em outra

doença.

Família Simuliidae Schiner

A família Simuliidae até o pre-

sente, conta com 8 gêneros, que

são: Simulium Latreille, 1802; Pro-

simulium Roubaud, 1906; Parasi-

mulium Malloch, 1914; Austrosi-

mulium Tonnoir, 1925; Cnephia

Enderlein, 1925; Gigantodax En-

derlein, 1925; Lutzsimulium Andr.

& Andr., 1946 e Gymnopais Stone,

1949. Dêstes, 2 ocorrem no Brasil,

o Simulium Latreille, 1802 o mais

antigo da família e o Lutzsimulium

Andr. & Andr., 1946, êste último

somente na parte meridional. Aqui

nos ocupamos somente do gênero

existente na região.

Gênero Simulium Latr.

A primeira forma conhecida do

gênero Simulium foi o imago, só

posteriormente se conhecendo a

larva, pupa e ovos.

Adulto — Cabeça livre, face in-

ferior breve, probóscida curta, for-

te e um pouco saliente; olhos re-

niformes e glabros; fronte sem oce-

los, sede de dimorfismo sexual evi-

dente; no macho, estreita e olhos

holópticos, na fêmea afastada e

olhos dicópticos; antena curta, ro-

busta com 11 artículos; palpo ma-

xilar com 5 segmentos, sendo o ba-

sal mais curto que os demais e o

terminal de comprimento variável,

no 3.° segmento localizando-se o ór-

gão sensorial de Lutz que apresen-

ta forma e tamanho variável. Es-

cudo abaulado sem sutura trans-

versal, recoberto de escamas e cer-

das que, às vêzes oferece ornamen-

tação e pruinosidade; escutelo glo-

boso semilunar. Pleuras e coxas um

tanto desenvolvidas com cerdas em

alguns escléritos cuja localização

entra na caracterização das espé-

9 — 37 145

130

Atas do Simpósio sobre a Biota Amazônica

cies. Pernas comparativamente ao

corpo, curtas, fortes um pouco

achatadas nos fêmures e tíbias;

metatarso alongado; os demais ar-

tículos tarsais curtos, o compri-

mento dos quatro reunidos é sem-

pre inferior ao comprimento do

metatarso; calcípala e pedisulco

presentes; unhas glabras ou den-

teadas, púlvilos rudimentares. Asa

hialina, comprida e larga sem cé-

lula basal e a nervura cubital on-

dulada sem bifurcação; as nervuras

das margens anteriores mais escu-

ras que as outras, estas às vêzes são

imperceptíveis; nervura costal, RI e

R 2 -j- 3 geralmente com espinhos

e cerdas, às vêzes na subcostal; álu-

las grandes com ângulo saliente;

esquâmula rudimentar. Balancins

salientes e inclinados sôbre o abdo-

me. Abdome com 8 segmentos vi-

síveis sendo o primeiro munido de

pêlos curtos ou longos, marginais;

órgão genital geralmente escon-

dido.

Pupa com um par de ganchos

terminais; segmentos abdominais

V, VI e VII com, no máximo 4 pa-

res de espinhos na face dorsal;

brânquias respiratórias de forma e

número de elementos variáveis.

Larva cilíndrica, caractística

acentuadamente delgada no meio,

na vista dorsal; antena com 4

segmentos; escovas bucais com

cêrca de 40 elementos; mandíbula

com 1 — 3 dentes; segmentos abdo-

minais I e II de menor diâmetro,

aumentando até alcançar a largu-

ra máxima do corpo nos segmen-

tos VI e VII, o diâmetro dêstes seg-

mentos é quase o dôbro dos das por-

ções I e II.

BIONOMIA

As larvas e pupas dêste gênero

são encontradas, como de resto as

demais espécies da família, em

águas agitadas ou então de certa

correnteza. Para postura dos ovos,

as fêmeas preferem as plantas her-

báceas à margem dos igarapés; os

galhos das árvores tombadas sôbre

êstes; as frestas das pedras nos lei-

tos mais ou menos secos, dos iga-

rapés, e, nos trechos onde a corren-

teza é mais intensa, porém não

muito forte, à espera das primeiras

águas, o que permite a eclosão das

larvas. Elas se alimentam de detri-

tos orgânicos e se fixam ao subs-