biodiversity

durrom

biodiversity

Membracidae: Wonder of terrestrial biodiversity.

Photos and text: Patrick Landmann/Lightmediation

Contact: Thierry Tinacci- Lightmediation Agency- +33 (0) 6 61 80 57 21

The astonishing biodiversity of

our planet is the result of a

perpetual and never-ended

evolution where the biological

species had to adapt to an often

hostile environment. The

complex relationship between

them and their habitat, the

pressure of the natural selection

and the spontaneous mutations,

between chance and necessity,

shaped living it in a multitude of

forms, which exceed our

imaginary. If all the mechanisms

of this evolutionary process are

not completely elucidated, it will

be sure that the dynamics of the

life has transformed, eliminated

and modified the species since

the beginning of time.


193-06: Bocydium globulare, fullface.


193-01/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-01: Cladonota latifrons.

An imitation which curiously points out a desiccated branchlet. / South America / Neotropical forest

193-03/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-03: Cladonota benitezi,

male. / South America / Neotropical forest

193-02/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-02: Cladonota benitezi,

female. Strong dimorphism with the male / South America / Neotropical forest

193-04/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-04: Smerdalea

imminens, fullface. / South America / Neotropical forest


193-14: Heteronotus maculatus, The hooked spines, horns and other points with which some species are provided in number wound the throat or are planted in the tissue which could then be infected and kill

later the animal consequently.


193-05/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-05: Smerdalea

imminens, female. / South America / Neotropical forest

193-07/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-07: Bocydium

globulare, in profile. / South America / Neotropical forest

193-06/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-06: Bocydium

globulare, fullface. / South America / Neotropical forest

193-08/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-08: Cladonota sp.

male. / South America / Neotropical forest


193-16: Umbelligerus peruviensis. The hooked spines, horns and other points with which some species are provided in number wound the throat or are planted in the tissue which could then be infected and

kill later the animal consequently.


193-09/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-09: Heteronotus

nigrogiganteus, in profile. The hooked spines, horns and other points with which some species are provided

in number wound the throat or are planted in the tissue which could then be infected and kill later the animal

193-11/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-11: Heteronotus

nigrogiganteus, from the top. The hooked spines, horns and other points with which some species are

provided in number wound the throat or are planted in the tissue which could then be infected and kill later

193-10/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-10: Heteronotus

nigrogiganteus, fullface. The hooked spines, horns and other points with which some species are provided

in number wound the throat or are planted in the tissue which could then be infected and kill later the animal

193-12/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-12: Heteronotus

delineatus. / South America / Neotropical forest


193-19: Heteronotus delineatus.


193-13/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-13: Heteronotus

delineatus, fullface. / South America / Neotropical forest

193-15/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-15: Heteronotus

albopunctatus. The hooked spines, horns and other points with which some species are provided in number

wound the throat or are planted in the tissue which could then be infected and kill later the animal

193-14/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-14: Heteronotus

maculatus, The hooked spines, horns and other points with which some species are provided in number

wound the throat or are planted in the tissue which could then be infected and kill later the animal

193-16/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-16: Umbelligerus

peruviensis. The hooked spines, horns and other points with which some species are provided in number

wound the throat or are planted in the tissue which could then be infected and kill later the animal


193-03: Cladonota benitezi, male.


193-17/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-17: Head of

Umbelligerus peruviensis. The hooked spines, horns and other points with which some species are

provided in number wound the throat or are planted in the tissue which could then be infected and kill later

193-19/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-19: Heteronotus

delineatus. / South America / Neotropical forest

193-18/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-18: Heteronotus

delineatus. / South America / Neotropical forest

193-20/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-20: Atypa bucktoni. /

South America / Neotropical forest


193-21/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-21: Anchistrotus

maculatus, has precuts at the base of its outgrowth which break or is detached when it is snapped up,

allowing the insect to escape. / South America / Neotropical forest

193-23/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-23: Lycoderes

gladiator, fullface. / South America / Neotropical forest

193-22/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-22: Anchistrotus

maculatus, fullface. / South America / Neotropical forest

193-24/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-24: Oeda inflata. /

South America / Neotropical forest


193-36: Enchophyllum cruentatum.


193-25/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-25: Cyphonia clavata.

While mating. / South America / Neotropical forest

193-27/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-27: Head of Stegaspis

fronditia, female. / South America / Neotropical forest

193-26/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-26: Stegaspis fronditia,

female. / South America / Neotropical forest

193-28/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-28: Stegaspis fronditia,

female from 3/4. / South America / Neotropical forest


193-29/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-29: Stegaspis fronditia,

female on a flower. / South America / Neotropical forest

193-31/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-31: Stegaspis fronditia

female. The excrements are rejected in form of small sweetened droplets, the honeydew. This very sticking

liquid is difficult to eliminate mechanically regarding the sedentary and the quasi-immobility of those

193-30/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-30: Stegaspis fronditia,

female with an ant Dolichoderus bispinosus. Mutualistic associations are very complex and multifactorial.

Mutualism are found between the two species only if the benefit is important for each one: for example,

193-32/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-32: Stictopelta squarus.

looks like the buds of the host plant amazingly. / South America / Neotropical forest


193-57: Certainly a larvae of Anchistrotus.


193-33/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-33: Head of Stictopelta

squarus. / South America / Neotropical forest

193-35/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-35: Membracis flaveola

in profil. / South America / Neotropical forest

193-34/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-34: Membracis

flaveola. / South America / Neotropical forest

193-36/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-36: Enchophyllum

cruentatum. / South America / Neotropical forest


193-37/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-37: Enchophyllum

cruentatum fullface. / South America / Neotropical forest

193-39/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-39: Lycoderes

fernandezi, female of its laying. / South America / Neotropical forest

193-38/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-38: Lycoderes

fernandezi, female of its laying. Gregarious they are scattered on the stem with their outgrowths pointed

towards outside make think of spines of certain shrubs. / South America / Neotropical forest

193-40/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-40: Tritropidia

bifenestrata, with a larvae. / South America / Neotropical forest


193-41/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-41: Larvae of

Tritropidia bifenestrata, babied by an ant Dolichoderus bispinosus. Mutualistic associations are very

complex and multifactorial. Mutualism are found between the two species only if the benefit is important for

193-43/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-43: Female of tritropidia

bifenestrata with egges, larvae and ants Dolichoderus bispinosus. Mutualistic associations are very complex

and multifactorial. Mutualism are found between the two species only if the benefit is important for each

193-42/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-42: Larvae an mature

Tritropidia bifenestrata on its laying. Mutualistic associations are very complex and multifactorial. Mutualism

are found between the two species only if the benefit is important for each one: for example, protection

193-44/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-44: Tritropidia

bifenestrata. When a young feels threatened, the vibrations so created are communicated within the group

of the larvae and the nymphs. Together, they tap the stem to alert the mothers who come to defend them by


193-01: Cladonota latifrons. An imitation which curiously points out a desiccated branchlet.


193-45/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-45: Oriola picta, female

on its laying. / South America / Neotropical forest

193-47/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-47: Gerridius fowleri,

female on its laying. / South America / Neotropical forest

193-46/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-46: Oriola picta, female

on its laying with young an ant Dolichoderus bispinosus. Mutualistic associations are very complex and

multifactorial. Mutualism are found between the two species only if the benefit is important for each one: for

193-48/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-48: Gerridius fowleri,

female on its laying with ant Dolichoderus bispinosus. Mutualistic associations are very complex and

multifactorial. Mutualism are found between the two species only if the benefit is important for each one: for


193-49/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-49: Lycoderes fabricii.

Its outgrowth pointed towards outside make think of spines of certain shrubs. / South America / Neotropical

forest

193-51/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-51: Enchenopa

albidorsa. Its outgrowth pointed towards outside make think of spines of certain shrubs. / South America /

Neotropical forest

193-50/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-50: Enchenopa

albidorsa, with a louse (in red). Its outgrowth pointed towards outside make think of spines of certain

shrubs. / South America / Neotropical forest

193-52/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-52: Enchenopa

gracillis. Its outgrowth pointed towards outside make think of spines of certain shrubs. / South America /

Neotropical forest


193-53/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-53: Nassunia binotata. /

South America / Neotropical forest

193-55/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-55: Stegaspis fronditia,

male. / South America / Neotropical forest

193-54/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-54: Bolbonota insignis,

This one looks like an bird excrement. A perfect imitation ! / South America / Neotropical forest

193-56/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-56: Tynelia pubescens.

/ South America / Neotropical forest


Membracidae:

Wonder of

terrestrial

biodiversity.

The astonishing biodiversity of our planet

is the result of a perpetual and

never-ended evolution where the

biological species had to adapt to an often

hostile environment. The complex

relationship between them and their

habitat, the pressure of the natural

selection and the spontaneous mutations,

between chance and necessity, shaped

living it in a multitude of forms, which

exceed our imaginary. If all the

mechanisms of this evolutionary process

are not completely elucidated, it will be

sure that the dynamics of the life has

transformed, eliminated and modified the

species since the beginning of time.

A real mother hen

Chef d'oeuvre of biological diversity,

Membracidae astonish us and let us

perplex because their forms are strange

and eccentric. The curious and impressive

expansions in front of their head or in

extension of their prothorax make real

alive sculptures of them. Tricks of nature

or mimicry, these insects somewhat

"baroques" surprise by the extravagance

and the great diversity of their outgrowths.

What were the environmental constraints

and pressures which in that way shaped

these odd expansions? The long path of

living molded these protuberances in

curious cuticular prolongations in front of

the head or starting from the first thoracic

segment (prothorax): spherical sizes,

curved or right spines, arabesques, horns,

roundnesses and more or less complex

structures form the Membracidae

exoskeleton. Their aspects are varied as

much: hairy to smooth, rough and

polygonal even reticulated or similar to

bark, dented, translucent or opaque,

colored or cryptic. These empty hulls, of

impressive sizes make the flight difficult

for some species, but Membracidae

remain very sharp in their jump. The

males can be very different from the

females regarding the morphology. After

mating, from which the positions are very

different according to the structure and the

form of the outgrowths, the female, at the

time of the laying, inserts directly in the

living tissue of their host plant, either a

single egg or plenty of them, or glue it on

the surface. Some species coat them with

a frothy substance which, while drying,

becomes hard. The very protective

females (they cover the eggs with their

body), will form groups by indifferently

mothering the larvae and the nymphs of

the ones and others until the adulthood,

which would lead us to believe that they

have a "maternal instinct".

Discreet, females are tented by ants

Because of their small size (approximately

1 cm length), Membracidae are often

unseen or very difficult to spot. Among the

2500 species which belong to the

Homoptera order, many, solitary and

discreet, remain badly known. Only those

are studied which are gregarious and

have mutualistic relations with ants.

Homoptera live on annual or perennial

plants and sucks the sap from which they

feed nutritive compounds thanks to a

complex digestive system. The

excrements are rejected in form of small

sweetened droplets, the honeydew. This

very sticking liquid is difficult to eliminate

mechanically regarding the sedentary and

the quasi-immobility of those species. It

isn't seldom to find an insect definitively

fixed on the host plant, killed

contaminated and invaded by moulds.

Their sweetened excrements are an

appreciable source of food for the arboreal

and opportunistic ants which intense

foraging activity facilitates the meetings

with the sedentary females which then

develop gregarious habits. Mutualistic

associations are very complex and

multifactorial. Mutualism are found

between the two species only if the benefit

is important for each one: for example,

protection (against the predatories) and

care (to inhibit parasitism, to avoid

diseases from fungus) in exchange of a

great production of honeydew. This

production is carried out when the ants

claim for and incite the evacuation of the

sweetened liquid in "stroking" the insect

with its antennas. The weak operating

cost for the ants, an easy access and a

short-haul until the place of harvest

increase association between the two

species. The host plant takes profit from

the protection thus brought to the

Membracidae by the ants because they

will push away all the specialized

plant-eating animals.

The predatory are wary about

The tropical forest is rich, plentiful of life.

From the litter untill the top of the trees,

life is omnipresent. The extraordinary

variety of the species is without end, the

multitude of the forms incredible. The

small insects of the Membracidae family

are one of the most significant

representatives. Neither aggressive, nor

equipped with natural defensive means

(darts, mandibles) so they probably

developed imitations in compensation that

could be at the origin of their outgrowths.

As these which looks like the buds of the

host plant amazingly, or those which,

scattered on the stem with their

outgrowths pointed towards outside make

think of spines of certain shrubs, or others

which prolongations remind fine

desiccated branchlets. First of all one

could think that this tiny world, if peaceful

and quiet. It is only one impression,

because some species share a very

advanced technique. Several types of

acoustic signal, inaudible sounds to

human, are transmitted thanks to the

vibrations made by tapping the legs on the

stem of the host plant for example. When

a young feels threatened, the vibrations

so created are communicated within the

group of the larvae and the nymphs.

Together, they tap the stem to alert the

mothers who come to defend them by

using the power of their posterior legs or

by beating their wings vigorously. Thus

they can push back predatory much more

bigger in size. Other sound vibrations are

used by the males which drum the plant

with their abdomen to attract females.

Predatories, like various Arthropods,

Arachnida and Hyménoptera (wasps) can

be pushed back by the mothers at first

sight so placid but foolhardy when their

offspring should be protected. Birds or

other predatories which which feed on

Membracidae are terribly wary. The

hooked spines, horns and other points

with which some species are provided in

number wound the throat or are planted in

the tissue which could then be infected

and then later kill the animal. To counter

the attacks of the animals, some species

of Membracidae have precuts at the base

of their outgrowths which break or are

detached when they are snapped up,

allowing the insect to escape.

These attractive insects, mini-monsters or

wonders of nature still keep very

mysterious. If one can easily observe most

common of them which live in partnerships

with the ants, how much are those, solitary

in the heights of the trees, that remain to

us unknown?

Membracidae, dazzling for the ones,

pushing

back for the others, but don't let indifferent

at all, are the witnesses of the immense

diversity of the species.


Captions

1 Cladonota latifrons. An imitation which

curiously points out a desiccated branchlet

2 Cladonota benitezi, female. Strong

dimorphism with the male (3).

3 Cladonota benitezi, male.

4 Smerdalea mminens, fullface.

5 Smerdalea imminens, female.

6 Bocydium globulare, fullface.

7 Bocydium globulare, in profile.

8 Cladonota sp. male.

9 Heteronotus nigrogiganteus, The

hooked spines, horns and other points

with which some species are provided in

number wound the throat or are planted in

the tissue which could then be infected

and kill later the animal consequently.

11 Heteronotus nigrogiganteus, from the

top.

12 Heteronotus delineatus.

13Heteronotus delineatus, fullface.

14 Heteronotus maculatus, see 9.

15 Heteronotus albopunctatus, see 9.

16 Umbelligerus peruviensis, see 9.

17 Head of Umbelligerus peruviensis.

18 Heteronotus delineatus.

19 Heteronotus delineatus, fullface.

20 Atypa bucktoni.

21 Anchistrotus maculatus, has precuts at

the base of its outgrowth which break or is

detached when it is snapped up, allowing

the insect to escape.

22 Anchistrotus maculatus, fullface.

23 Lycoderes gladiator, fullface.

24 Oeda inflata.

25 Cyphonia clavata. While mating.

26 Stegaspis fronditia, female.

27 Head of Stegaspis fronditia, female.

28 Stegaspis fronditia, female from 3/4.

29 Stegaspis fronditia, female on a

flower.

30 Stegaspis fronditia, female with an ant

Dolichoderus bispinosus, see 41.

31 Stegaspis fronditia female. The

excrements are rejected in form of small

sweetened droplets, the honeydew. This

very sticking liquid is difficult to eliminate

mechanically regarding the sedentary and

the quasi-immobility of those species. It

isn't seldom to find an insect definitively

fixed on the host plant, killed

contaminated and invaded by moulds.

32 Stictopelta squarus. looks like the buds

of the host plant amazingly.

33 Head of Stictopelta squarus.

34 Membracis flaveola.

35 Head of Membracis flaveola.

36 Enchophyllum cruentatum.

37 Enchophyllum cruentatum, de face.

38 Lycoderes fernandezi, female of its

laying. Gregarious they are scattered on

the stem with their outgrowths pointed

towards outside make think of spines of

certain shrubs.

39 Idem Lycoderes fernandezi, female of

its laying.

40 Tritropidia bifenestrata. with a larvae.

41 Larvae of Tritropidia bifenestrata,

babied by an ant Dolichoderus bispinosus.

Mutualistic associations are very complex

and multifactorial. Mutualism are found

between the two species only if the benefit

is important for each one: for example,

protection (against the predatories) and

care (to inhibit parasitism, to avoid

diseases from fungus) in exchange of a

great production of honeydew. This

production is carried out when the ants

claim for and incite the evacuation of the

sweetened liquid in "stroking" the insect

with its antennas. The weak operating

cost for the ants, an easy access and a

short-haul until the place of harvest

increase association between the two

species. The host plant takes profit from

the protection thus brought to the

Membracidae by the ants because they

will push away all the specialized

plant-eating animals.

42 Larvae an mature Tritropidia

bifenestrata on its laying, see 41.

43 Female of tritropidia bifenestrata with

egges, larvae and ants Dolichoderus

bispinosus, see 41.

44 Tritropidia bifenestrata. When a young

feels threatened, the vibrations so created

are communicated within the group of the

larvae and the nymphs. Together, they tap

the stem to alert the mothers who come to

defend them by using the power of their

posterior legs or by beating their wings

vigorously.

45 Oriola picta, female on its laying.

46 Oriola picta, female on its laying, seer

41.

47 Gerridius fowleri, female on its laying.

48 Gerridius fowleri, female on its laying

with ant Dolichoderus bispinosus.

49 Lycoderes fabricii.

50 Enchenopa albidorsa, with a louse (in

red).

51 Enchenopa albidorsa, see 38.

52 Enchenopa gracillis, see 38.

53 Nassunia binotata.

54 Bolbonota insignis, This one looks like

an bird excrement. A perfect imitation !

55 Stegaspis fronditia, male.

56 Tynelia pubescens.

57 Certainly a larvae of Anchistrotus.

For more informations :

Hôlldobler, B. & E.O. Wilson. 1990. The

ants. The Belknap Press of Harvard

University Press, Cambridge,

Massachusetts.

(Gullan, P.J. and Kosztarab, M. -1997-

Adaptations in scale insects. Annual

Review of Entomology 42: 23-50).

Rex Cocroft, The inside story of insect

song, American Museum of Natural

History, October 1999

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