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.

Membracidae: Wonder of terrestrial biodiversity.

Photos and text: Patrick Landmann/Lightmediation
Contact: Thierry Tinacci- Lightmediation Agency- +33 (0) 6 61 80 57 21
193-06: Bocydium globulare, fullface.
193-01/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-01: Cladonota latifrons. 193-02/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-02: Cladonota benitezi,
An imitation which curiously points out a desiccated branchlet. / South America / Neotropical forest female. Strong dimorphism with the male / South America / Neotropical forest

193-03/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-03: Cladonota benitezi, 193-04/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-04: Smerdalea
male. / South America / Neotropical forest 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 193-06/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-06: Bocydium
imminens, female. / South America / Neotropical forest globulare, fullface. / South America / Neotropical forest

193-07/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-07: Bocydium 193-08/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-08: Cladonota sp.
globulare, in profile. / South America / Neotropical forest 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 193-10/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-10: Heteronotus
nigrogiganteus, in profile. The hooked spines, horns and other points with which some species are provided 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 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 193-12/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-12: Heteronotus
nigrogiganteus, from the top. The hooked spines, horns and other points with which some species are delineatus. / South America / Neotropical forest
provided in number wound the throat or are planted in the tissue which could then be infected and kill later
193-19: Heteronotus delineatus.
193-13/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-13: Heteronotus 193-14/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-14: Heteronotus
delineatus, fullface. / South America / Neotropical forest 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

193-15/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-15: Heteronotus 193-16/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-16: Umbelligerus
albopunctatus. The hooked spines, horns and other points with which some species are provided in number 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 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 193-18/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-18: Heteronotus
Umbelligerus peruviensis. The hooked spines, horns and other points with which some species are delineatus. / South America / Neotropical forest
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 193-20/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-20: Atypa bucktoni. /
delineatus. / South America / Neotropical forest South America / Neotropical forest
193-21/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-21: Anchistrotus 193-22/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-22: Anchistrotus
maculatus, has precuts at the base of its outgrowth which break or is detached when it is snapped up, maculatus, fullface. / South America / Neotropical forest
allowing the insect to escape. / South America / Neotropical forest

193-23/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-23: Lycoderes 193-24/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-24: Oeda inflata. /
gladiator, fullface. / South America / Neotropical forest South America / Neotropical forest
193-36: Enchophyllum cruentatum.
193-25/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-25: Cyphonia clavata. 193-26/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-26: Stegaspis fronditia,
While mating. / South America / Neotropical forest female. / South America / Neotropical forest

193-27/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-27: Head of Stegaspis 193-28/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-28: Stegaspis fronditia,
fronditia, female. / South America / Neotropical forest female from 3/4. / South America / Neotropical forest
193-29/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-29: Stegaspis fronditia, 193-30/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-30: Stegaspis fronditia,
female on a flower. / South America / Neotropical forest 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,

193-31/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-31: Stegaspis fronditia 193-32/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-32: Stictopelta squarus.
female. The excrements are rejected in form of small sweetened droplets, the honeydew. This very sticking looks like the buds of the host plant amazingly. / South America / Neotropical forest
liquid is difficult to eliminate mechanically regarding the sedentary and the quasi-immobility of those
193-57: Certainly a larvae of Anchistrotus.
193-33/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-33: Head of Stictopelta 193-34/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-34: Membracis
squarus. / South America / Neotropical forest flaveola. / South America / Neotropical forest

193-35/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-35: Membracis flaveola 193-36/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-36: Enchophyllum
in profil. / South America / Neotropical forest cruentatum. / South America / Neotropical forest
193-37/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-37: Enchophyllum 193-38/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-38: Lycoderes
cruentatum fullface. / South America / Neotropical forest 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

193-39/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-39: Lycoderes 193-40/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-40: Tritropidia
fernandezi, female of its laying. / South America / Neotropical forest bifenestrata, with a larvae. / South America / Neotropical forest
193-41/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-41: Larvae of 193-42/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-42: Larvae an mature
Tritropidia bifenestrata, babied by an ant Dolichoderus bispinosus. Mutualistic associations are very Tritropidia bifenestrata on its laying. Mutualistic associations are very complex and multifactorial. Mutualism
complex and multifactorial. Mutualism are found between the two species only if the benefit is important for are found between the two species only if the benefit is important for each one: for example, protection

193-43/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-43: Female of tritropidia 193-44/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-44: Tritropidia
bifenestrata with egges, larvae and ants Dolichoderus bispinosus. Mutualistic associations are very complex bifenestrata. When a young feels threatened, the vibrations so created are communicated within the group
and multifactorial. Mutualism are found between the two species only if the benefit is important for each 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 193-46/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-46: Oriola picta, female
on its laying. / South America / Neotropical forest 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

193-47/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-47: Gerridius fowleri, 193-48/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-48: Gerridius fowleri,
female on its laying. / South America / Neotropical forest 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. 193-50/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-50: Enchenopa
Its outgrowth pointed towards outside make think of spines of certain shrubs. / South America / Neotropical albidorsa, with a louse (in red). Its outgrowth pointed towards outside make think of spines of certain
forest shrubs. / South America / Neotropical forest

193-51/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-51: Enchenopa 193-52/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-52: Enchenopa
albidorsa. Its outgrowth pointed towards outside make think of spines of certain shrubs. / South America / gracillis. Its outgrowth pointed towards outside make think of spines of certain shrubs. / South America /
Neotropical forest Neotropical forest
193-53/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-53: Nassunia binotata. / 193-54/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-54: Bolbonota insignis,
South America / Neotropical forest This one looks like an bird excrement. A perfect imitation ! / South America / Neotropical forest

193-55/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-55: Stegaspis fronditia, 193-56/©Patrick Landmann/Lightmediation/Wonder of terrestrial biodiversity / 193-56: Tynelia pubescens.
male. / South America / Neotropical forest / 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 45 Oriola picta, female on its laying.
towards outside make think of spines of
certain shrubs. 46 Oriola picta, female on its laying, seer
41.
39 Idem Lycoderes fernandezi, female of
its laying. 47 Gerridius fowleri, female on its laying.
40 Tritropidia bifenestrata. with a larvae. 48 Gerridius fowleri, female on its laying
with ant Dolichoderus bispinosus.
41 Larvae of Tritropidia bifenestrata,
babied by an ant Dolichoderus bispinosus. 49 Lycoderes fabricii.
Mutualistic associations are very complex
and multifactorial. Mutualism are found 50 Enchenopa albidorsa, with a louse (in
between the two species only if the benefit red).
is important for each one: for example,
protection (against the predatories) and 51 Enchenopa albidorsa, see 38.
care (to inhibit parasitism, to avoid
diseases from fungus) in exchange of a 52 Enchenopa gracillis, see 38.
great production of honeydew. This
production is carried out when the ants 53 Nassunia binotata.
claim for and incite the evacuation of the
sweetened liquid in "stroking" the insect 54 Bolbonota insignis, This one looks like
with its antennas. The weak operating an bird excrement. A perfect imitation !
cost for the ants, an easy access and a
short-haul until the place of harvest 55 Stegaspis fronditia, male.
increase association between the two
species. The host plant takes profit from 56 Tynelia pubescens.
the protection thus brought to the
Membracidae by the ants because they 57 Certainly a larvae of Anchistrotus.
will push away all the specialized
plant-eating animals. For more informations :
Hôlldobler, B. & E.O. Wilson. 1990. The
42 Larvae an mature Tritropidia ants. The Belknap Press of Harvard
bifenestrata on its laying, see 41. University Press, Cambridge,
Massachusetts.
43 Female of tritropidia bifenestrata with
egges, larvae and ants Dolichoderus (Gullan, P.J. and Kosztarab, M. -1997-
bispinosus, see 41. Adaptations in scale insects. Annual
Review of Entomology 42: 23-50).
44 Tritropidia bifenestrata. When a young
feels threatened, the vibrations so created Rex Cocroft, The inside story of insect
are communicated within the group of the song, American Museum of Natural
larvae and the nymphs. Together, they tap History, October 1999
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.