Teletusa Limpida (SIGNORET)

Teletusa limpida (SIGNORET): a Neotropical proconiine leafhopper that mimics megachilid bees (Hymenoptera: Apoidea), with notes on Batesian mimicry in the subfamily Cicadellinae (Hemiptera: Cicadellidae)

G. MEJDALANI, D.M. TAKIYA, M. FELIX, P. CEOTTO & D. YANEGA

Abstract

Morphological comparisons and field and M. neoxanthoptera. Yellow areas on studies carried out in the Brazilian State of the leafhopper abdomen resemble the Minas Gerais suggested that the proconi- metasomal scopal hairs or the pollen ine leafhopper Teletusa limpida (SlG- carried on the scopa of the bees. The NORET) is a Batesian mimic of bees of the present case of mimicry is compared with family Megachilidae. This leafhopper the two other known cases of mimicry in exhibits the following mimicry-related leafhoppers of the subfamily Cicadellinae characteristics: (1) body densely covered (genera Propetes WALKER and Lissoscarta by conspicuous hairs, especially developed STAL), as well as with similar cases in the on the face and mesoscutellum, (2) disc of Auchenorrhyncha. The appearance of frons forming an approximately right angle mimicry in the Cicadellinae is discussed in with the longitudinal axis of body, (3) a phylogenetic context. fore- and hindwings almost entirely hyaline, (4) abdomen short and expanded laterally, and (5) overall coloration dark Key words brown or black with yellow markings. These features are remarkably similar to Proconiini, Cicadellini, mimetic spe- those of four megachilid species that are cies, morphology, phylogeny, Megachili- sympatric with T. limpida in Minas Gerais dae, Vespidae. (Anthodioctes megachiloides (HOLMBERG), Hypanthidiodes subarenarium (SCHWARZ), Hypanthidiodes musciforme (SCHROTTKY), and Megachile meoxanthoptera COCKE- RELL. Specimens of T. limpida are espe- Denisia 04. cially similar to those of A. megachiloides zugleich Kataloge des OÖ. Landesmuseums. Neue Folge Nr. 176 (2002), 215-224

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Introduction the constriction at the base of the abdomen (data based so far only on field observations of Batesian mimicry in the subfamily L. vespiformis (FABRICIUS) and L. beckeri Cicadellinae MEJDALANI &. FELIX). This behavior, in which Two cases of Batesian mimicry in leafhop- the mimic suddenly exposes characteristics pers of the subfamily Cicadellinae have been that are similar to those of its model, is called reported in the literature. BOULARD (1978) "ostensible mimicry" (BOULARD 1978). It has and MEJDALANI & FELIX (1997) described not been observed in any other known leaf- morphological and behavioral features of the hopper genus.

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cicadelline genus Lissoscarta Stäl that enable TAK1YA et al. (1999) described morpho- Figs 1 and 2. its seven known species to mimic epiponine logical features of the proconiine genus Propetes (Fig. 1) The cicadelline Batesian mimic Lissoscarta beckeri MEJDALANI & FEUX wasps (Vespidae: Polistinae) (Figs 1 and 2). WALKER related with the mimicry of epiponi- (female) and (Fig. 2) its epiponine The abdomen in Lissoscarta is strongly con- ne wasps. Specimens of the two known spe- wasp model Agelaia fulvofasciata cies of Propetes, as those of Lissoscarta, have stricted at the base, the fore- and hindwings (DEGEER), body in dorsal view hyaline and elongate forewings and the abdo- (northern Brazil; from MEJDALANI & are elongate and hyaline, and the color pat- men is constricted basally. According to FEUX 1997). Mimic and model are tern is remarkably similar to that of many epi- approximately the same size. TAKIYA et al. (1999), the males of P. schmidti ponine wasps (crown, pronotum, and meso- MELICHAR are mostly black with small yellow notum yellow with dark maculae and stripes; areas and stripes, while in the females these abdominal terga yellowish-brown with a yellow markings are distinctly larger (Figs 3 transverse yellow stripe on their posterior and 5). The different color patterns of males margins). When threatened, Lissoscarta leaf- and females suggested the existence of a dual, hoppers spread their wings in a similar way to sex-limited Batesian mimicry. In this kind of that of a wasp in the resting position, showing mimicry each sex of the mimic has its own

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Figs 3 and 4. (Fig. 3) The proconiine Batesian mimic Propetes schmidti MEUCHAR (male) and (Fig. 4) its epiponine wasp model Polybia rejecta (FABRIOUS), body in dorsal view (southeastern Brazil; from TAKIYA et al. 1999). Mimic and model are approximately the same size.

Figs 5 and 6. (Fig. 5) The proconiine Batesian mimic Propetes schmidti MEUCHAR (female) and (Fig. 6) its epiponine wasp model Mischocyttarus ypiranguensis da FONSECA, body in dorsal view (southeastern Brazil; from TAKIYA et al. 1999). Mimic and model are approximately the same size.

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models (VANE-WRIGHT 1976) (Figs 4 and 6). spectively by MEJDALANI & FELIX (1997) and This is the only known case of dual-mimicry TAKIYA et al. (1999) as belonging to class VI in the Auchenorrhyncha. According to VANE- (antergic defensive). According to VANE- WR1GHT (1976), this kind of mimicry occurs WR1GHT (1976), in this class the mimic simu- lates an organism in some way repellent to the rarely in Lepidoptera and more commonly in operator, so avoiding attention it would other- Hymenoptera (e.g., EVANS 1968). wise receive. The presence of the mimic is a Using VANE-WRIGHT'S (1976) terminolo- disadvantage to the model. Class VI includes gy and analytic schemes, the mimicry cases of H.W. Bates' original formulation of mimicry Lissoscarta and Propetes were considered re- (Fig. 10).

Figs 7-9. (Fig. 7) The proconiine Batesian mimic Teletusa limpida (SIGNORET) (male) and (Fig. 9) its megachiline bee model Megachile neoxanthoptera COCKEREU, body in dorsal view (southeastern Brazil). Mimic and model are approximately the same size. (Fig. 8) T. limpida. anterior portion of body in lateral view.

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The proconiine genus Teletusa Field studies carried out in the Brazilian DISTANT State of Minas Gerais by one of the authors (DY) suggested that T. limpida could be a YOUNG (1968) recognized a single valid species in the Neotropical genus Teletusa, mimic of long-tongued solitary bees of the T. limpida (SlGNORET), with six junior syno- family Megachilidae (leafcutting bees). The nyms. He examined specimens of Teletusa morphological analysis of T. limpida leafhop- from Amazonas in Brazil to Bolivia and to pers and megachilid bees from the locality Paraguay and Argentina. The results of a phy- (Belo Horizonte) studied by DY confirmed the logenetic analysis of the proconiine genera existence of such a mimicry relationship. The

Bees (Megachilinae) — - Teletusa Bees (Megachilinae)—Bees (Megachilinae) Wasps (Polistinae)—Propetes, Lissoscarta Wasps (Polistinae) — Wasps (Polistinae)

R Predator Predator CLASS VI CLASS IA 10 11

with the exposed posterior meron (MEJDA- bees examined, which belong to the subfami- Figs 10 and 11. (Fig. 10) Scheme showing the LANI 2000) indicated that Teletusa is the sister ly Megachilinae, are Anthodioctes megachilo- known Batesian mimicry (class VI) group of a clade formed by the genera Abana ides (HOLMBERG), Hypanthidiodes (Moureanth- cases in the leafhopper subfamily DISTANT, Acrobelus STÄL, Deselvana YOUNG, idium) subarenarium (SCHWARZ), Hypanthid- Cicadellinae (based on VANE-WRIGHT 1976). In class VI, antergic defensive and Rhaphirrhinus LAPORTE. The clade in- iodes {Saranthidium) musciforme (SCHROTTKY) mimicry, the presence of the mimic is cluding the latter four genera and Teletusa is (tribe Anthidiini), and Megachile (Ptilosar- a disadvantage to the model. supported by two apomorphic features: (1) oides) neoxanthoptera COCKERELL (tribe Mega- (Fig. 11) Müllerian mimicry (class IA) cases treated in the present study posterior region of crown with an elevated chilini). We provide below an account of the (based on VANE-WRIGHT 1976). In class area between the ocelli and (2) apex of aede- characteristics of T. limpida that enable it to IA, synergic warning mimicry, the agus with a pair of lateral lobes. Teletusa can mimic the megachilid bees. These character- model and mimic cooperate to produce a common defense signal that adver- be distinguished from the remaining genera of istics are compared with those of the proposed tises their dangerous, nauseous or the clade by the following apomorphic models. The present case of mimicry is com- otherwise repellent features to the operator. S-), model (signal transmit- characteristics: (1) body densely covered by pared with the two other known cases of ter); S2, mimic (signal transmitter); conspicuous hairs, especially developed on mimicry in leafhoppers of the subfamily Cica- R, operator (signal receiver); the face and mesoscutellum (Fig. 8), (2) disc dellinae, as well as with similar cases in the +, advantageous interactive role; -, disadvantageous interactive role. of frons forming an approximately right angle Auchenorrhyncha. with the longitudinal axis of body (Fig. 8), (3) mesoscutellum swollen (Fig. 8), (4) forewings Material and methods almost entirely hyaline (Fig. 7), and (5) abdomen short and expanded laterally (Fig. 7). The specimens of T. limpida examined, YOUNG (1968) observed that Teletusa stands which were determined by one of the authors well apart from other genera in the tribe Pro- (GM), are deposited in the following collec- coniini in its very pubescent mesoscutellum tions: Departamento de Entomologia, Museu and lower portion of the face, in the very Nacional, Universidade Federal do Rio de short pygofer of the male, and in the Janeiro (MNRJ; Rio de Janeiro), Departa- broad, flattened, short abdomen. mento de Zoologia, Museu Paraense Emflio

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Goeldi (MPEG; Belem), Departamento de (5) Megachile neoxanthoptera. Brazil: State Zoologia, Instituto de Biologia, Universidade of Minas Gerais: 5 males and 3 females, Belo Federal do Rio de Janeiro (DZRJ; Rio de Horizonte Municipality (UFMG); 1 male, Janeiro), and Departamento de Zoologia, Ponte Nova Municipality (UFMG). Setor de Ciencias Biolögicas, Universidade The techniques for preparation of the Federal do Parana (DZUP; Curitiba). The male genital structures, which were dissected megachilid specimens studied are deposited in to confirm the identification of T. limpida sen- the Departamento de Zoologia, Instituto de su YOUNG (1968: 164, Fig. 151), follow those Ciencias Biolögicas, Universidade Federal de of OMAN (1949). The dissected parts are Minas Gerais (UFMG; Belo Horizonte). They stored in microvials with glycerin. The leaf- were determined by the Drs. F. A. Silveira hopper morphological terminology adopted (UFMG) or D. Urban (DZUP). The sexes and herein follows mainly MEJDALANI (1998, 2000), locality data of all studied specimens are as fol- while the bee morphological terminology follows: lows mainly MITCHELL (1980) and MiCHENER (1) Teletusa limpida. Brazil: State of Ama- (1944). The body length (in mm) of leafhop- zonas: 1 male, Säo Gabriel da Cachoeira pers and bees was taken from the apex of head Municipality (MNRJ); 1 male, Maturacä [Säo to the apex of abdomen in dorsal view. The Gabriel da Cachoeira Municipality] (MPEG); mimicry concepts, terminology, and analytic 1 male and 3 females, Manaus Municipality schemes used herein follow mainly the work of (MNRJ); State of Para: 1 male, Serra Norte VANE-WRIGHT (1976). [Parauapebas Municipality] (MPEG); State of Rondonia: 1 male, Vilhena Municipality Results (MNRJ); State of Mato Grosso: 1 male, Cha- pada dos Guimaräes Municipality (MPEG); Megachilid bees State of Goiäs: 8 males, Campinas [Goiänia Males and females of the four studied bee Municipality] (MNRJ); State of Minas Gerais: species vary in length, respectively, from 5.6 to 2 males, Belo Horizonte Municipality 8.1 mm and from 6.0 to 9.9 mm (see details on (MNRJ); State of Säo Paulo: 3 males, Ilha Table 1). The body in these bees has a mostly Seca [Andradina Municipality] (MNRJ); 1 black, nonmetallic ground color, and is thick- male and 1 female, Bebedouro Municipality ly beset with hairs (more conspicuously de- (DZUP). Peru: Department of Loreto: 1 male, veloped in Megachile neoxanthoptera) on the AguasCalientes[?](DZRJ). head, thorax, and abdomen (Fig. 9). The con- (2) Anthodioctes megachiloides. Brazil: State tour of the face, in lateral view, forms an of Minas Gerais: 3 males and 5 females, Belo approximately right angle with the longitudi- Horizonte Municipality (UFMG). nal axis of body. The fore- and hindwings are (3) Hypanthidiodes subarenarium. Brazil: mostly hyaline. The abdomen (metasoma) has State of Minas Gerais: 1 male and 1 female, an ellipsoidal form, or is somewhat rounded Belo Horizonte Municipality (UFMG); 2 (males of M. neoxanthoptera). Terga on the males and 1 female, Catas Altas Municipality posterior half of the metasoma are covered by (UFMG). yellow pubescence (M. neoxanthoptera), or (4) Hypanthidiodes musciforme. Brazil: possess conspicuous transverse yellow stripes State of Minas Gerais: 1 male, Belo Horizonte (Anthodioctes megachiloides, Hypanthidiodes Municipality (UFMG); 2 males and 1 female, subarenarium, and H. musciforme). The pollen- Table 1. Sahara Municipality (UFMG). collecting scopa is on the metasoma, not on Body length (in mm) of males and females of the leafhopper Teletusa limpida (SIGNORET) and of the megachi- Males Females lid bees Anthodioctes megachiloides Teletusa limpida 6.2-7.2 (n= 13) 7.1-8.7 (n = 4) (HOLMBERG), Hypanthidiodes subaren- Anthodioctes meaachiloides 5.6-6.2 (n = 3) 6.8-7.9 (n = 5) arium (SCHWARZ), Hypanthidiodes musciforme (SCHROTTKY), and Megachile Hypanthidiodes subarenarium 6.4-6.7 (n = 3) 6.0-6.3 (n = 2) neoxanthoptera COCKERELL. The number Hypanthidiodes musciforme 6.3-7.5 (n = 3) 6.7 (n = 1) (n) of specimens measured is given inside parentheses. Megachile neoxanthoptera 6.5-8.1 (n = 6) 9.4-9.9 (n = 3)

220 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at the hindlegs as in most bees. It occupies most independently in the clade that includes the of the metasomal sternum, being formed by genus Propetes, the wasp-mimic proconiine long, robust, densely distributed hairs. The (TAKIYA et al. 1999, MEJDALANI 2000) (Figs 3 sting is well developed. and 5). Interestingly, this feature also arose independently in the wasp-mimic cicadelline Teletusa limpida Lissoscana (BOULARD 1978, MEJDALANI & The above-mentioned characteristics of FELIX 1997, FELIX 1999, MEJDALANI 2000) the megachilid bees are apparently mimicked (Fig. 1). The hyaline hindwings, which are by Teletusa limpida (Figs 7-9). Males and also mimicry-related, are common in the females of this leafhopper vary in length, Cicadellidae, being a plesiomorphic trait in respectively, from 6.2 to 7.2 mm and from 7.1 the hymenopteran-mimic leafhoppers. The to 8.7 mm (Table 1). The body has a dark fore- and hindwings of Teletusa, Propetes, and brown or black ground color and is remark- Lissoscana (Figs 1, 3, 5, and 7) clearly resem- ably covered by hairs (Figs 7 and 8), being ble those of bees or wasps (Figs 2, 4, 6, and 9). very similar to that of the bees, especially In addition to this important role, the hyaline A. megachibides and M. neoxanthoptera (Fig. 9). aspect possibly allows an easier visualization The hairs are strongly developed on the by the predators (operators) of the leafhopper mesoscutellum and face; the contour of the abdomen, which is short and expanded lat- latter, in lateral view, also forms an approxi- erally (Teletusa, Fig. 7) or has a distinct con- mately right angle with the longitudinal axis striction {Lissoscana and Propetes, Figs 1, 3, of body (Fig. 8). The abdominal sterna, latero- and 5). tergites, terga vii and viii, and the pygofer Leafhoppers of the genus Teletusa, as well (tergum ix) are variably marked with yellow. as those of the genera Propetes and Lissoscarta, These yellow areas resemble the scopal hairs are very poorly represented in collections. or the pollen carried on the scopa, as well as They are apparently rare insects, an aspect color markings or hairs on other metasomal that supports the hypothesis of Batesian areas of the bees. The fore- and hindwings are, mimicry (class VI, antergic defensive, of as in the proposed models, mostly hyaline. VANE-WRIGHT 1976) (Fig. 10). The predators The abdomen is short, broad, and slightly (operators) from which the Batesian mimicry flattened dorsoventrally, clearly resembling in Propetes and Lissoscarta affords protection the metasoma of the bees. Its lateral areas are are unknown. Thus, it was not possible to visible at the sides when the wings are at rest determine whether such mimicry cases are dis- position. junct (mimic, model, and operator are different species) or bipolar (model and operator Discussion are the same) (MEJDALANI & FELIX 1997, TAKIYA et al. 1999). In the Teletusa case the The aforementioned autapomorphic fea- predators are also unknown. This case, how- tures of Teletusa, which clearly distinguish it ever, is certainly disjunct, since the model from other related proconiine genera (YOUNG bees are not predators. 1968, MEJDALANI 2000), are apparently mimicry-related, with the single exception of At least two other cases of class VI the expanded mesoscutellum. The hairy body, mimicry have been reported in the Auche- perpendicular frons, short and laterally ex- norrhyncha, both in the planthopper family panded abdomen, and hyaline forewings (as well Fulgoridae. HOGUE (1984) suggested that as hindwings), along with the dark brown or "lantern bugs", Fulgora spp., avoid predation black overall coloration with yellow markings, by mimicking lizards. He observed that indi- give T. limpida (Figs 7 and 8) an appearance viduals of Fulgora typically rest during the day remarkably similar to the megachilids studied, on the trunks of large trees. They position especially A. megachiloides and M. neoxantho- themselves vertically, the head with its great ptera (Fig. 9). The hairs and the peculiar aspect anterior protuberance uppermost and elevated of the frons and abdomen are unique within at an angle away from the substratum. This the tribe Proconiini (MEJDALANI 2000). The posturing is similar to that assumed by certain hyaline forewings, on the other hand, arose species of arboreal lizards, e.g., Plica plica (LlN-

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NAEUS) (Iguanidae). HOGUE (1984) described because the conspicuous spines on its prono- similarities in the size, overall form, and gen- tum would keep predators away. Thus, the ves- eral color pattern of "lantern bugs" and lizards, poid mimetic signal would be reinforced. He as well as remarkable points of resemblance in did not mention whether H. flawhneatus is a their heads (posture, shape, and color Batesian or Müllerian mimic of ants. Personal markings) (Fig. 12). ZOLNEROWICH (1992) observations by DY revealed that Heteronotus described a nymph of Amycle sp. that mimics treehoppers are particularly similar to eumenine vespids. These wasps are typically black jumping spiders (Salticidae). He observed that and yellow, with a doubly-constricted metaso- the nymph has four smooth, polished dark ma. The membracids are similarly marked, with a greatly elongated pronotum with two strong posterior constrictions, giving them the same size and proportions as the eumenine wasps. VANE-WRIGHT (1976) observed that the models in class VI mimicry are often members of Müllerian mimicry groups (class IA, synergic warning) (Fig. 11). According to him, in class IA the model and mimic co-operate to produce a common defense signal that adver- tises their dangerous, nauseous or otherwise repellent features to the operator. This is the classic situation originally described by F. Mül- ler. One example of such Müllerian groups, or rings, are the epiponine wasps that are mimicked by Lissoscarta and Propetes leafhop- Fig. 12. areas on its metathorax and hindwing pads pers (RICHARDS & RICHARDS 1951, BOULARD The „lantern bug" Fulgora sp. (Ful- goridae), head, thorax, and part of that resemble the anterior eyes of jumping 1978, MEJDALANI & FELIX 1997, TAKIYA et al. wings in lateral view (northern spiders. The middle- and hindlegs, which 1999) (Figs 2 and 6). Our conclusion that Brazil). According to HOGUE (1984), project posteriorly and have flattened femora members of two tribes of the subfamily Mega- „lantern bugs" are possibly Batesi- and tibiae, are moved in an up and down an mimics of arboreal lizards. The chilinae could be models for T. limpida is ap- mimics are similar to the proposed manner similar to those of salticid legs and parently in accordance with the observation models in the size, overall form, pedipalpi during courtship or aggression of VANE-WRIGHT (1976) that the models in and general color pattern, inclu- displays. In the cases reported by HOCUE class VI often belong to Müllerian groups. As ding remarkable points of resemblance in their heads (posture, sha- (1984) and ZOLNEROWICH (1992) the model described before, the body in the four studied pe, and color markings). and operator were considered the same bees has a very similar color pattern. It is (bipolar). mostly black, nonmetallic, with the tergal Other cases of mimicry in the Auchenorr- posterior half of the metasoma mainly yellow hyncha, which so far have not been classified (Fig. 9). Interestingly, the yellow metasomal area in M. neoxanthoptera (Megachilini) and according to the concepts of VANE-WRIGHT (1976), occur in the treehopper genus Heter- those in A. megachiloides, H. subarenarium, onotus LAPORTE (Membracidae) (BOULARD and H. musciforme (Anthidiini) are not 1999, DY, personal observations). BOULARD homologous. In the former species the yellow (1999) reported the occurrence of mimicry of area is formed by pubescence, while in the lat- ants and wasps by species of Heteronotus. He ter three it is formed by transverse, colored observed that individuals of H. flavolineatus integumental stripes. Such non-homologous LAPORTE are apparently ant-mimics. They are conditions suggest that these bees belong to a usually found with these hymenopterans, Müllerian ring. Their sting, along with the being sedentary and gregarious. On the other hairy body, would teach predators to keep hand, individuals of H. abcisus WALKER are away. This suggestion of a Müllerian ring is solitary and more active fliers, being possibly corroborated by field observations in Belo wasp-mimics. According to him, the latter Horizonte (Minas Gerais). In this area species could belong to a Müllerian ring, T. limpida appears to belong to a diverse group

222 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at of palatable and unpalatable mimics that malskomplexe der Zikade zurückzuführen: includes wasps (Sphecidae and Vespidae), (1) eine auffällige und dichte Behaarung des bees (Apidae, Colletidae, and Megachilidae), Körpers, insbesondere im Gesicht und am moths (Arctiidae), flies (Syrphidae), and Mesoscutellum, (2) die Platte der Frons steht beetles (Cerambycidae) (DY, personal obser- im rechten Winkel zur Körperlängsachse, vations). (3) Vorder- und Hinterflügel sind fast zur Gänze transparent, (4) das Abdomen ist kurz Acknowledgments und lateral verbreitert, und (5) die Färbung ist dunkelbraun bis schwarz mit gelben Flecken. We are greatly indebted to F. A. Silveira Diese Merkmale sehen jenen von vier Mega- (UFMG) for providing invaluable information chiliden-Arten, die mit T. Umpida in Minas and literature on the family Megachilidae and Gerais sympatrisch vorkommen {Anthodioctes for making available for study bee specimens megachiloides (HOLMBERG), Hypanthidiodes under his care. The following persons lent subarenarium (SCHWARZ), H. musciforme specimens ofTektusa: R.R. Cavichioli (DZUP), (SCHROTTKY) und Megachik neoxanthoptera N. Ferreira-Jr. (DZRJ), A.Y. Harada (MPEG), COCKERELL), sehr ähnlich. Individuen von and J.L. Nessimian (DZRJ). The manuscript T. Umpida sehen vor allem A. megachiloides benefited from the useful comments of A.C.R. und M. neoxanthoptera sehr ähnlich. Die gel- Alves, A.L. Carvalho, C.H. Dietrich, R. Raki- ben Flecken am Abdomen der Zikade ähneln tov, and F.A. Silveira. The habitus drawings of den Haaren der Scopa der Bienen oder Pollen, Lissoscarta beckeri and Agelaia fulvofasciata auf der Scopa transportiert wird. Die hier were inked by M.F. Pessoa, and those of Prop- beschriebene Mimikry wird mit zwei anderen etes schmidti, Tektusa Umpida, Mischocyttarus Fällen von Mimikry bei Zikaden (Gattungen ypiranguensis, Polybia rejecta, and Megachik Propetes WALKER und Lissoscarta STÄL; beides neoxanthoptera were inked by L.A.A. Costa. Cicadellinae) verglichen. Ab-schließend wer- The photograph of Fulgora sp. was taken by den phylogenetische Aspekte des Auftretens M. J. G. Hopkins. Fellowships from the von Mimikry bei Zikaden diskutiert. following Brazilian agencies are greatly acknowledged: Coordenacäo de Aperfeigoamento de Pessoal de Nfvel Superior (CAPES) to PC, References MF, and DMT, Conselho Nacional de Desen- volvimento Cientifico e Tecnologico (CNPq) BOULARD M. (1978): Premier cas de „mimetisme to DMT and DY, and Fundacäo de Amparo ä ostensible" chez les Homopteres Auchenorhyn- Pesquisa do Estado de Säo Paulo (FAPESP) to ques (Insecta). — C. R. Acad. Sc. Paris 287 (D): 1389-1391. GM. This study was supported in part by Uni- versidade Federal do Rio de Janeiro (UFRJ) BOULARD M. (1999): Contributions a I'entomologie generate et appliquee. 2. Cicadaires (Rhynchota and Fundacäo Universitäria Jose Bonifacio Auchenorhyncha). Deuxieme partie: Membra- (FUJB). S. A. Vanin was the adviser of GM coidea. I.- Notes et documents. — EPHE, Biol. during his D.Sc. studies in the Instituto de Bio- Evol. Insec. 11/12: 141-182, pls. 1-9. ciencias da Universidade de Säo Paulo (USP). EVANS J.W. (1968): Studies on Neotropical Pompilidae The advisership and encouragement provided (Hymenoptera) IV. Examples of dual sex-limited mimicry in Chirodamus. — Psyche 75: 1-22. by S. A. Vanin are greatly acknowledged. FELIX M. (1999): Anälise filogenetica do genero Lissoscarta STAL, 1869 (Hemiptera: Cicadellidae: Cicadellinae).— M. Sc. Dissertation, Universidade Zusammenfassung Federal do Rio de Janeiro, xiii + 117 pp. HOGUE C.L. (1984): Observations on the plant hosts and possible mimicry models of "Lantern bugs" Morphologische Befunde und Freilandstu- (Fulgora spp.) (Homoptera: Fulgoridae). — Rev- dien in Minas Gerais (Brasilien) zeigen, dass ta Biol. Trop. 32: 145-150.

Gestalt und Färbung der Zwergzikade Tektusa MEJDALANI G. (1998): Morfologia externa dos Cicadel- Umpida (SlGNORET) aus der Unterfamilie linae (Homoptera: Cicadellidae): comparacäo Cicadellinae (Tribus Procimiini) als Batesische entre Versigonalia ruficauda (WALKER) (Cicadelli- ni) e Tretogonia cribrata MEUCHAR (Proconiini), Mimikry der Bienenfamilie Megachilidae zu com notas sobre outras especies e anälise da interpretieren sind. Dies ist auf fünf Merk- terminologia. — Revta Bras. Zool. 15: 451-544.

MEJDALANI G. (2000): Morfologia externa e anälise cladistica dos generös de Proconiini com o mero posterior exposto (Hemiptera, Cicadellidae, Cicadellinae). — D. Sc. Thesis, Universidade de Säo Paulo, Säo Paulo, xiii + 221 pp.

MEJDALANI G. & M. FEUX (1997): A new species of the Neotropical genus Lissoscarta STAL (Homoptera: Cicadellidae: Cicadellinae) that mimics wasps. — Proc. Ent. Soc. Wash. 99: 156-161.

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TAKIYA D.M., MEJDALANI G. S M. FELIX (1999): Dual- mimicry of wasps by the Neotropical leafhopper Propetes schmidti MELICHAR with a description of its female (Hemiptera: Cicadellidae: Cicadelli- nae). — Proc. Ent. Soc. Wash. 101: 722-728. Addresses of the authors: VANE-WRIGHT R.I. (1976): A unified classification of mimetic resemblances. — Biol. J. Linn. Soc. Prof. Dr. Gabriel MEJDALANI, Lond. 8: 25-56. Paula CEOTTO, and Märcio FELIX YOUNG D.A. (1968): Taxonomic study of the Cicadelli- Departamento de Entomologia nae (Homoptera: Cicadellidae). Part 1, Proconi- ini. — Bull. U. S. Natn. Mus. 261: 1-287. Museu Nacional ZOLNEROWICH G. (1992): A unique Amycle nymph Universidade Federal do Rio de (Homoptera: Fulgoridae) that mimics jumping Janeiro spiders (Araneae: Salticidae). — J. N. Y. Ent. Soc. 100: 498-502. Quinta da Boa Vista, Säo Cristöväo 20940-040 Rio de Janeiro, RJ, Brasil E-mail: [email protected]

Daniela M. TAKIYA Departamento de Zoologia Universidade Federal do Rio de Janeiro Rio de Janeiro, RJ, Brasil Present address: Center for Biodiversity Illinois Natural History Survey 607 East Peabody Drive Champaign, IL 61820, USA

Dr. Douglas YANEGA Department of Entomology University of California Riverside, CA 92521-0314, USA

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