Rev. Biol. Trop., 32(1): 145-150, 1984

Observations on the plant hosts and possible mimicry models of "Lantern Bugs" ( spp.) (Homoptera : )

Charles L. Hogue Natural History Museum of. Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007 U.S.A.

(Received for publication January 4, 1984)

Abstract:From the author's field obse.rvations and literature records, neotropical "Lantern Bugs" (Fulgora spp.) are now known to use tree species of 5 families as resting, and presumably, feeding hosts: Simarouba- . ceae- Aublet; Simaba versicolor St. Hilaire ;Fabaceae Hy menaea oblongifolia (var. palustris (Ducke) Lee & Langenheim, Hy menaea coubaril Linnaeus;Myroxylon balsamum (Linnaeus) ; Rutaceae·-Zantho­ xylum sp. ; Lecvthidaceae-Lecythis sp,; Vochysiaceae- Vochysia tucanorum Martius. Most of these are pro­ ducers of resins or noxious chemicals which may be sequestered by the bugs. These trees are also fr equented by various arboreal lizards which are good candidates for models for mimicry by the bugs because of the reptilian-like appearance of the latter's large head protuberance and other structural and color features. The resemblance of Linnaeus to Plica plica (Linnaeus) in the Iquitos area of Peru is especially close and this is postulated as the model in that area.

The very large homopterans known popu­ the host trees of these curious bugs and add larly in English as "Lantem Bugs", "Peanut sorne cautious speculations on the mime tic Headed Bugs", or "Alligator Headed Bugs" function of their body fo rm, particularly that (Fulgora spp.) are common and widely dis­ of the unique cephalic projection. tributed through the fo rests of the Neotropical Region. Specimens often are seen on tree HOSTS trunks and frequentIy fly to artificial lights at night. The 11 species recognized in the genus To one searching fo r these in the (Metcalf 1947; Brailovsky and Beutelspacher Peruvian rain forest, it soon becomes apparent 1978) have long attracted the attention of that they occur only on a few kinds of large laymen and entomologists alike, because of trees, from among the many available. They their great size and especially the grotesque , are seen on the boles but may also choose inflated prolongation of the head. Their other parts upon which to rest and presumably conspicuousness and bizarre appearance have fe ed. The bugs are a tourist attraction and precipitated considerable note in the literature specimens command a price as souvenirs. Thus, and led to fanciful stories of alleged venomous­ the natives, many of whom have spent their ness, sound producing abilities, resemblance to entire lives hunting and roaming in these crocodilians, and luminescence. However, little regions, are able to take the visitor directIy to reliable scientific information on their biology such trees. With the help of local guides, I was has been published. able to locate several host specimens and collect In the years 1979 to 1983, I spent a few data on the insecto weeks each June-July in the forests near Iqui­ In the vicinity of Iquitos, the trees most tos, Peru, where I observed a number of often hosting F. laternaria are Simarouba specimens of F. la ternaria Linnaeus under amara Aublet, a species in the familiy Sima­ natural conditions and interviewed several roubaceae, known thereabouts as "marupá" persons familiar with local natural history and Hy menaea oblongifo lia var. palustris regarding the habits of the species. From these (Ducke) Lee & Langenheim, in the Fabaceae, field contacts (and review of the literature) I called 10calIy "azucar-huayo". To date, I have am able to offer the following information on not personally seen Fulgora on trees of these

145 146 REVISTA DE BIOLOGIA TROPICAL

species that 1 have visited, but am assured by fe rred to as My roxylon balsamum (Linnaeus) three reliable informants that the insects Harms, known also in the Spanish American habitually visit them at other times of the vernacular as "Balsam of Pe ru " , or "quino­ year. One guide reported dozens of individuals quino" on the trunks of the former tree at a single Select plants of five different families are time. At a locality near the confluence of the now known to serve as hosts of Fulgora and Rios Napo and Yagua, 1 examined one marupá one wonders about sorne common factor which, although not at the time harboring that makes them attractive to or of service any Fulgora , was infested with over 20 me­ to these insects . They either produce and dium-sized fulgorids of another type, Lystra concentrate resins (Hymenaea, My roxylon, lanata Linnaeus, and numerous large zygopine Vochysia), oils (Lecythis, eucalyptus?) or (Ceratosomus) and choline (Homalinotus) bitter substances in their sap (­ weevils, indicating sorne unexplained attractive­ pers. obs., Simarouba, Simaba-Porter 1973), ness to these insects. The tree was emitting i.e. possibly generating allelopathic chemicals considerable amounts of resin from bark not identifiable at this time (See Uphof, 1968 wounds and fr om an apparently diseased for general chemical products of these trees). trunk near the ground. The presence of such allelopaths is implied by "Guapinol", which is Hy menaea coubari! these categories of substances which often have Linnaeus, a relative of azucar-huayo, is a well toxic or noxious qualities and which might be known host of Fulgora sp. in Costa Rican transferred to their succivorous parasites. dry forests (Janzen and Hogue, 1983). This Sesquiterpines, for example, known plant tree does not grow in the wet forests near allomones, have been isolated from Hy menaea Iquitos although other Hy menaea attract these (Lee and Langenheim, 1975) and terpenoids homopterans there . Allen M. Young (pers. are known to be sequestered by insects and comm.) reports this plant as especially at­ used defensively by sorne insects (Blum, 1981: tractive to in Central America. 440-44 1). A third confirmed host at Yanamono, 35 The existence of a volatile defensive che m­ km northeast of Iquitos, is a single ical is suggested by DeVries' observation (in Zanthoxylum sp. from which 1 have personally lanzen and Hogue, 1983) that Fu lgora, when collected F. laternaáa. The species, atypical persistently and sufficiently molested, may leap of the genus in its lack of trunk spines, remains into flight, ernitting a fe tid odor. No glands unidentified (Gentry per. comm.). specifically for the production of noxious Simarouba amara is given as a definitehost odors are known to exist on the bug's body; by Moss (in Poulton, 1932) in Brazil and such volatiles could reside in the body's cover­ Bolivia and Alwyn H. Gentry (pers. comm.) ing of wax [ Note : Complex a1cohols are pre­ from a single tree near 1 auneche, western sent in the copious wax secretions of Attacus Ecuador. Gentry noted other numerous large atlas Linnaeus, a silk moth which feeds on the fulgorids (Lystra ?) also in association with "Tree of Heaven", Ailan th us altissima (Miller), this tree. Another member of the same family, another member of the Simaroubaceae (J ones Simaba versicolor St . Hilaire, called "pau et al. , 1982)] parahyba" locally was considered by Branner lean H. Langehheim (pers. comm.),suggests (1885) to be a tree well known to be fre­ members of the genus Copaifera, including quented by Fulgora in eastern Brazil, a fact the so-called "Diesel Fuel Tree" or "copahiba", noted much earlier by Chabrillac (1859, as "Pa­ C. multijuga Hayne, in Amazonia as additional rahiba"). Branner mentioned, incidentally, that candidates for the list of Fulgora hosts. eucalyptus was considered by many people to have an attraction for the bugs in this region. MIMICRY Only one other account (Lenko and Papavero 1979) lists hosts for these bugs; in Brazil these are "caixetta", "pau de tucano", "resinera", Individuals of Fulgora typically rest during Vochysia tucanorum Martius (Vochysiaceae); the day on the trunks of the aforementioned "supucaias", Lecythis spp. (Lecythidaceae); trees. They position themselves vertically, and "pau de oleo", My rospermum the head with its great anterior protuberance erythroxylum Allemao (Fabaceae), now re- uppermost and elevated at an angle away HOGUE: Possible mimicry models of Lantern Bugs 147

Fig. 1. Possible mimicry of a lizard by a homopterous insecto Fig. la. Plica plica, Sauria. Fig. lb. Fulgora latemaria, Insecta. Photographs of live specimens from 35 km northeast Iquitos, Peru, by C. 1. Hogue. Scale lines equal 25 mm each.

from the substratum. This posturing is While observing F. laternaria on the trunk of similar to that assumed by certain species of the Zan th oxylum at Yanamono on one occa­ arboreal iguanid lizards, whose appearance in sion 1 was startled to realize that on the same this way and otherwise suggests a resting bug. tree a pair of medium-sized specimens of the 148 REVISTA DE BIOLOGIA TROPICAL lizard Plica plica (Linnaeus), from the base of Since noting these similarities in the field the tail, across the back and along the elongate, between P. plica and Fulgara laternaria, I have slightly reared head, much resembled resting searched among museum collections and the Fulgara laternaria. Soon after I was able to literature for other potential models among examine in hand a female lizard of this species the Sauria, and even tree frogs and arboreal collected from another tree (Fig. la) and found snakes that might prove to be more precise numerous detailed points of resemblance models,. incIuding ones that might serve as between elements of its structure and that of models for species of Fulgara showing markings the bug (Fig. 1 b). and protuberance shapes different from F. These elements are : (1) size . Fulgara are la ternaria. It appears from these comparisons extremely large members of their family, that the bugs may be mimicking a variety of having at least 3 to 4 times the body bulk of arboreal iguanid species which inhabit surfaces any other neotropical species, and approxi­ of large tree boles andfor branches. The lizards mating the body size of these lizards. Their (Dixon and Soini, 1975; Duellman , 1978; apparent size is increased by the expanse of Etheridge, 1979; Vanzolini, 1972) are most the wings, which when folded, cover an area likely to be found in the genera Analis, roughly equal to a medium-sized lizard's Enyaliades, Enyalius, Palychms, Enyaliasaums, abdomen; (2) overall form, omitting the lizard's Uranascadan , and even juvenile Ig uana. Less tail, which is very slender and therefore not likely but possible models may also exist among obvious, its wide, compressed abdomen and Basiliscus and Carytaphanes, and possibly sorne narrowed head is very much like the bug's speeies of Gekkonidae. The "leaf nosed" or total form viewed from aboye or obliquely; "proboseis" anoles, A. punctatus and relatives (3) general color pattern. (a) Both bug and (Williams, 1965), may be specific models for F. lizard are mottled green and black. (b) AIso, slender nosed bug types like cracadilus lucile ra the bug's forelegs have bands similar to those Brailovsky and Beutlespacher or F. on the lizard's legs; (4) head posture. As already Germar. stated, heads of both the lizards and bugs are These ideas must be regarded as speculative held obliquely erect; (5) head shape. The heads at this time. Field studies of antipredator of the two trace very nearly the same behavior as well as geographic correlations between various models and Fulgara speeies outline in profile; both are (a) moderately • elongate and cylindrical, with a (b) dorsal are required before this mimetic association prominence just anterior to the middle, (c) can be verified. There seems no doubt, however , arched anterodorsal surface, (d) bluntly pointed that species of Fulgara are mimicking a reptili­ apex and (e) slightly convex undersides; (6) an rather than amphibian form, as evidenced head markings. Structure on the lizard's head by the unmistakable scale-like markings on the are simuláted by markings on the bug's head lower sides of the head protuberance. In spite in the following manner: (a) dark, circular of the amazing parallels in head form, the field surroundíng the bug's eye equals the suggestion that the model is to be fo und lizard's tympanum or anterior dark part of the among the Crocodilians (Poulton, 1924; Hin­ nuchal collar, (b) dark field with anterior emar­ ton, 1977) is untenable, since the bugs are gination beneath the dorsal convexity on the strictly arboreal. That it is among the Sauria bug simulates the lizard's highlighted eye, (c) rather than Serpentes, rests on the a�umed small black spot near the apex of the bug's pro­ simulation of the tympanum by the circum­ tuberance equals the lizard's nostril (d) a do u­ ocular macula and is strengthened by the other ble row of squarich maculae along the side points of similarity listed aboye as well as strict subventrally on the bug simulates the lizard's syntopy. mouth bordered aboye and below with Yet sorne likeness to certain snakes must be dark-edgad labial scales and (e) a finely granular admitted still, namely the lateral square mottling on the underside appearing very much maculae to the labial scales and pits of boids, like the guIar region of the reptile. and in sorne Fu lgara species, a black spot . Incongruence of the strongIy arching line between the false eye and nostril to the loreal just anterior to the bug's eye protuberance fo ssa of arboreal pit vipers ( Bathraps bi­ may be a vestigial state common to virtually lineatus, B. schlegeli and B. venezuelensis). all fulgoroids with a porrect cephalic swelling. Of possible significance also is the practice HOGUE: Possible mimicry models of Lantern Bugs 149

of Brazilian Indians to call these insects in Branner, J. C. 1885. The reputation of the lantern Tupi "jaquirana-boia" or "snake ", from fly. Amer. Nat.,19' 835-838. iaquirana = cicada + mboi = snake (Lenko and Chabrillac. M. t's59. [ Communication at society Papavero, 1979) nec "jacareenam-boya" = "alligator snake" as mistranslated by Francis meetingl , Bull. Soco Entomolg. France, Ser 3 7:cii.

Walker (opinion of Poulton, 1924 : xlvi). The Dixon, J. R., & P. Soini. 1975. The reptiles of the 's partial resemblance to the anterior upper Amazon Basin. Iquitos Region, Peru. 1. part of a snake, however, might explain the Lizards and Amphisbaenians. Contrib. Biol. Geol. ubiquitous fe ar of lantern bugs throughout Milwaukee Pub. Mus., 5 1-5 8. South America. Duellman, W. E. 1978. The biology of an equatorial Using Vane Wright's (1976) terminology herpetofauna in Amazonian Ecuador; Univ. Kansas and analytic schemes, I would define this case Mus. Nat. Hist. Mise. Pub! ., 65: 1-352. of mimicry as Class VI, antergic defensive, in which the model and the operator (as a Etherigde, R. 1970. A review of the South American iguanid lizard genus Plica. ,BulI. British Museum predator) are the same (bipolar, Sl+ R). It is (Nat. Hist.), Zool., 19: 237-256. c1earIy Batesian and most c10sely parallels an artifice fo und in the lepidopteran family Hinton, H. E. 1977. Mimicry jJrovides information Brassolidae, employing hindwing markings abou t the perceptual capacities of predators. Folia similar to the silhouettes of tree frogs and Entomol. Mexicana, 37 : 19-29. anoles which inhabit tree trunks where the Janzen, D. H., & C. 1. Hogue. 1983. Fulgara laternaria butterflies often rest (Stradling, 1976). The (Machaca, Peanut-head Bug, Lantern Fly), p. most proximate and likely oí" the bug's 726-727, In D. H. Janzen (ed.). Costa Rican Natural predators logically would be these insectivorous History (University of Chicago Press), xi, 816 p. lizards, from whose predation they would Jones, C.G., A. M. Young, T. H. Jones, & M. S. Blum. escape by resembling them. 1982. Chemistry and possible roles of cuticular a1cohols of the larval atlas moth. Comp. Biochem. ACKNOWLEDGMENTS Physio!., 73B: 797-801.

Lee, Y. & J. H. Langenheim. 1975. Systematics ofthe would like to thank the following indi­ genus Hy menaea L. (Leguminosae, Caesalpinoi­ viduals for invaluable assitance in this study : deae, Detarieae). Univ. California Pub!. Bot., 69: Drs. Roy W. McDiarmid, U.S. Fish and Wildlife 1-109. Service ; John W. Wright, Robert L. Bezy and Lenko, K. & N. Papavero, 1979. Insetos no folclore. John Karges of my own institution, in herpeto­ Conselho Estadual de Artes e Ciencias Humanos, logical matters; Drs. Jean H. Langenheim , Uni­ Sao Paulo, 518 p. versity of California, Santa Cruz, Al Gentry, Missouri Botanical Garden, and Mildred Ma­ Metcalf, Z. P. 1947. FasciCle IV, Fulgoroidea.Part. 9, thias, University of California, Los Angeles, fo r Fulgoridae. General Catalogue of the . Smith College,Northhampton,Massachusetts. 276 p. identification of botanical materials. My infor­ mants were Alfredo Chavez, Walter Navarro and Antero Cabral, all employees of Exploraciones Porter, D. M. 1973, Family 90. Simaroubaceae, Part Amazónicas S. A. Financial support was VI. 60: 23-39. In R. E. Woodson, Jr. & R. W. Schery and collaborators, Flora of Panama. Ann. provided by the Natural History Museum of Missouri Bot. Gard. Los Angeles County Foundation and Univer­ sity of California at Los Angeles Extensibn, Potilton, E. B. 1924. The terrifying appearance of Department of Biological and Physical Sciences. Laternaria (Fulgoridae) fo unded on the most prominent features of the alligator. Trans. En­ tomol. Soco London. xliii-xlix. LITERATURE CITED Poulton, E. B. 1932. The alligator-like hea.d and thorax of the tropical American Laternaria laternaria, 1. Blum, M. S. 1981. Chemical defenses of . (Fulgoridae, Homoptera). Proc. Entomol. Soco Academic Press. New York, xii, 562 p. London. 7: 68-70. Brailovsky, H. & C.R. Beutelspacher. 1978. Una nueva especie de Fulgara Linneo (Homoptera: Fulgori­ Stradling, D. J. 1976. The nature of the mimetic dae) de México. Ann. Inst. Biol., Univ. Nac. Aut. patterns of the brassolid genera, Ca liga and Ery­ México, Ser. Zool., 49: 175-1 82. phamis. Ecol. Entomol., 1: 135-138. 150 REVISTA DE BIOLOGIA TROPICAL

Uphof, J.C.T. 1968. Dictionary of economic plants, Vanzolini, P. E. 1972. Miscellaneous notes on the 2d ed. Verlag von J. Cramer, Lehre, 591 p. ecology of sorne Brazilian lizards (Sauria) Papo Avu!. Zoo!. sao Paulo, 26 : 83-115.

Vane-Wright, R. 1. 1976. A unified c1assification of Williams, E. E. 1965. South American Anolis (Sauria, mimetic - resemblances. Bio!. J. Linnean Soc., 8: Iguanidae) : Two new species of the Punctatus 25-56. Group. Breviora, 233: 1-15.