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
Transcript
The astonishing biodiversity ofour planet is the result of aperpetual and never-endedevolution where the biologicalspecies had to adapt to an oftenhostile environment. Thecomplex relationship betweenthem and their habitat, thepressure of the natural selectionand the spontaneous mutations,between chance and necessity,shaped living it in a multitude offorms, which exceed ourimaginary. If all the mechanismsof this evolutionary process arenot completely elucidated, it willbe sure that the dynamics of thelife has transformed, eliminatedand modified the species sincethe beginning of time.
Membracidae: Wonder of terrestrial biodiversity.Photos and text: Patrick Landmann/Lightmediation
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 killlater the animal consequently.
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 andkill later the animal consequently.
The astonishing biodiversity of our planetis the result of a perpetual andnever-ended evolution where thebiological species had to adapt to an oftenhostile environment. The complexrelationship between them and theirhabitat, the pressure of the naturalselection and the spontaneous mutations,between chance and necessity, shapedliving it in a multitude of forms, whichexceed our imaginary. If all themechanisms of this evolutionary processare not completely elucidated, it will besure that the dynamics of the life hastransformed, eliminated and modified thespecies since the beginning of time.
A real mother henChef d'oeuvre of biological diversity,Membracidae astonish us and let usperplex because their forms are strangeand eccentric. The curious and impressiveexpansions in front of their head or inextension of their prothorax make realalive sculptures of them. Tricks of natureor mimicry, these insects somewhat"baroques" surprise by the extravaganceand the great diversity of their outgrowths.What were the environmental constraintsand pressures which in that way shapedthese odd expansions? The long path ofliving molded these protuberances incurious cuticular prolongations in front ofthe head or starting from the first thoracicsegment (prothorax): spherical sizes,curved or right spines, arabesques, horns,roundnesses and more or less complexstructures form the Membracidaeexoskeleton. Their aspects are varied asmuch: hairy to smooth, rough and
polygonal even reticulated or similar tobark, dented, translucent or opaque,colored or cryptic. These empty hulls, ofimpressive sizes make the flight difficultfor some species, but Membracidaeremain very sharp in their jump. Themales can be very different from thefemales regarding the morphology. Aftermating, from which the positions are verydifferent according to the structure and theform of the outgrowths, the female, at thetime of the laying, inserts directly in theliving tissue of their host plant, either asingle egg or plenty of them, or glue it onthe surface. Some species coat them witha frothy substance which, while drying,becomes hard. The very protectivefemales (they cover the eggs with theirbody), will form groups by indifferentlymothering the larvae and the nymphs ofthe ones and others until the adulthood,which would lead us to believe that theyhave a "maternal instinct".
Discreet, females are tented by ants Because of their small size (approximately1 cm length), Membracidae are oftenunseen or very difficult to spot. Among the2500 species which belong to theHomoptera order, many, solitary anddiscreet, remain badly known. Only thoseare studied which are gregarious andhave mutualistic relations with ants.Homoptera live on annual or perennialplants and sucks the sap from which theyfeed nutritive compounds thanks to acomplex digestive system. Theexcrements are rejected in form of smallsweetened droplets, the honeydew. Thisvery sticking liquid is difficult to eliminatemechanically regarding the sedentary andthe quasi-immobility of those species. Itisn't seldom to find an insect definitivelyfixed on the host plant, killedcontaminated and invaded by moulds.Their sweetened excrements are anappreciable source of food for the arborealand opportunistic ants which intenseforaging activity facilitates the meetings
with the sedentary females which thendevelop gregarious habits. Mutualisticassociations are very complex andmultifactorial. Mutualism are foundbetween the two species only if the benefitis important for each one: for example,protection (against the predatories) andcare (to inhibit parasitism, to avoiddiseases from fungus) in exchange of agreat production of honeydew. Thisproduction is carried out when the antsclaim for and incite the evacuation of thesweetened liquid in "stroking" the insectwith its antennas. The weak operatingcost for the ants, an easy access and ashort-haul until the place of harvestincrease association between the twospecies. The host plant takes profit fromthe protection thus brought to theMembracidae by the ants because theywill push away all the specializedplant-eating animals.
The predatory are wary aboutThe tropical forest is rich, plentiful of life.From the litter untill the top of the trees,life is omnipresent. The extraordinaryvariety of the species is without end, themultitude of the forms incredible. Thesmall insects of the Membracidae familyare one of the most significantrepresentatives. Neither aggressive, norequipped with natural defensive means(darts, mandibles) so they probablydeveloped imitations in compensation thatcould be at the origin of their outgrowths.As these which looks like the buds of thehost plant amazingly, or those which,scattered on the stem with theiroutgrowths pointed towards outside makethink of spines of certain shrubs, or otherswhich prolongations remind finedesiccated branchlets. First of all onecould think that this tiny world, if peacefuland quiet. It is only one impression,because some species share a veryadvanced technique. Several types ofacoustic signal, inaudible sounds tohuman, are transmitted thanks to the
vibrations made by tapping the legs on thestem of the host plant for example. Whena young feels threatened, the vibrationsso created are communicated within thegroup of the larvae and the nymphs.Together, they tap the stem to alert themothers who come to defend them byusing the power of their posterior legs orby beating their wings vigorously. Thusthey can push back predatory much morebigger in size. Other sound vibrations areused by the males which drum the plantwith their abdomen to attract females.Predatories, like various Arthropods,Arachnida and Hyménoptera (wasps) canbe pushed back by the mothers at firstsight so placid but foolhardy when theiroffspring should be protected. Birds orother predatories which which feed onMembracidae are terribly wary. Thehooked spines, horns and other pointswith which some species are provided innumber wound the throat or are planted inthe tissue which could then be infectedand then later kill the animal. To counterthe attacks of the animals, some speciesof Membracidae have precuts at the baseof their outgrowths which break or aredetached when they are snapped up,allowing the insect to escape.
These attractive insects, mini-monsters orwonders of nature still keep verymysterious. If one can easily observe mostcommon of them which live in partnershipswith the ants, how much are those, solitaryin the heights of the trees, that remain tous unknown? Membracidae, dazzling for the ones,pushing back for the others, but don't let indifferentat all, are the witnesses of the immensediversity of the species.
Captions
1 Cladonota latifrons. An imitation whichcuriously points out a desiccated branchlet
2 Cladonota benitezi, female. Strongdimorphism 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, Thehooked spines, horns and other pointswith which some species are provided innumber wound the throat or are planted inthe tissue which could then be infectedand kill later the animal consequently.
11 Heteronotus nigrogiganteus, from thetop.
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 atthe base of its outgrowth which break or isdetached when it is snapped up, allowingthe 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 aflower.
30 Stegaspis fronditia, female with an antDolichoderus bispinosus, see 41.
31 Stegaspis fronditia female. Theexcrements are rejected in form of smallsweetened droplets, the honeydew. Thisvery sticking liquid is difficult to eliminatemechanically regarding the sedentary andthe quasi-immobility of those species. Itisn't seldom to find an insect definitivelyfixed on the host plant, killedcontaminated and invaded by moulds.
32 Stictopelta squarus. looks like the budsof 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 itslaying. Gregarious they are scattered onthe stem with their outgrowths pointedtowards outside make think of spines ofcertain shrubs.
39 Idem Lycoderes fernandezi, female ofits laying.
40 Tritropidia bifenestrata. with a larvae.
41 Larvae of Tritropidia bifenestrata,babied by an ant Dolichoderus bispinosus.Mutualistic associations are very complexand multifactorial. Mutualism are foundbetween the two species only if the benefitis important for each one: for example,protection (against the predatories) andcare (to inhibit parasitism, to avoiddiseases from fungus) in exchange of agreat production of honeydew. Thisproduction is carried out when the antsclaim for and incite the evacuation of thesweetened liquid in "stroking" the insectwith its antennas. The weak operatingcost for the ants, an easy access and ashort-haul until the place of harvestincrease association between the twospecies. The host plant takes profit fromthe protection thus brought to theMembracidae by the ants because theywill push away all the specializedplant-eating animals.
42 Larvae an mature Tritropidiabifenestrata on its laying, see 41.
43 Female of tritropidia bifenestrata withegges, larvae and ants Dolichoderusbispinosus, see 41.
44 Tritropidia bifenestrata. When a young feels threatened, the vibrations so createdare communicated within the group of thelarvae and the nymphs. Together, they tapthe stem to alert the mothers who come todefend them by using the power of theirposterior legs or by beating their wingsvigorously.
45 Oriola picta, female on its laying.
46 Oriola picta, female on its laying, seer41.
47 Gerridius fowleri, female on its laying.
48 Gerridius fowleri, female on its layingwith ant Dolichoderus bispinosus.
49 Lycoderes fabricii.
50 Enchenopa albidorsa, with a louse (inred).
51 Enchenopa albidorsa, see 38.
52 Enchenopa gracillis, see 38.
53 Nassunia binotata.
54 Bolbonota insignis, This one looks likean 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. Theants. The Belknap Press of HarvardUniversity Press, Cambridge,Massachusetts.
(Gullan, P.J. and Kosztarab, M. -1997-Adaptations in scale insects. AnnualReview of Entomology 42: 23-50).
Rex Cocroft, The inside story of insectsong, American Museum of NaturalHistory, October 1999
![Neotropical cervidology_12[1]](https://static.documents.pub/doc/80x56/547f4ab5b37959a22b8b56e0/neotropical-cervidology121.jpg)
