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Zeitschrift/Journal: Denisia

Jahr/Year: 2002

Band/Volume: 0004

Autor(en)/Author(s): Dietrich Christian O., Dietrich Christian O.

Artikel/Article: Evolution of (Insecta, ) 155-170 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Evolution of Cicadomorpha (Insecta, Hemiptera)

C.H. DIETRICH

Abstract

Cicadomorpha (Cicadoidea, Cerco- and tribe. The origins of some - poidea and ) are one of the group taxa may also have coincided with dominant groups of plant-feeding , shifts in feeding or courtship strategies, or as evidenced by their extraordinary diver- the colonization of novel habitats (e.g., sity and ubiquity in habitats ranging from grasslands, deserts). The origins of genera tropical rainforest to tundra. Improve- and , in many cases, can be attribu- ments on our knowledge of the phylogeny ted to shifts in habitat and host plant asso- of these insects, based on cladistic analysis ciation, as well as smaller scale biogeogra- of morphological and molecular data and phic vicariance. Many aspects of cicado- study of the record, provide the morphan evolution remain poorly under- opportunity to examine the possible fac- stood. These include phenomena such as tors that led to their diversification. Fac- the coexistence of many closely related tors influencing early divergences among species on the same host plant and the major lineages apparently included shifts diversity of bizarre pronotal modifications in life history strategies, including a tran- found among Membracidae. Such questi- sition from subterranean or cryptic to ons are best addressed by further ecologi- arboreal nymphal stage, shifts in feeding cal and behavioral study, as well as phylo- strategy (xylem to phloem or parenchy- genetic analysis. ma), and the acquisition of various mor- phological adaptations (crypsis, jumping Key words: , spittlebug, leafhop- hind legs, specialized grooming structu- per, , adaptation, evolution. res). Although most modern families pre- sently are distributed worldwide, global plate tectonics undoubtedly contributed to Denisia 04, diversification at the level of subfamily zugleich Kataloge des OÖ. Landesmuseums, Neue Folge Nr. 176 (2002), 155-170

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Introduction This paper reviews the current state of knowledge of cicadomorphan phylogenetic Cicadomorpha, comprising modern cica- relationships and suggests some possible fac- das (Cicadoidea), spittlebugs (Cercopoidea), tors that contributed to the origin and diversi- and (Membracoidea, fication of the various lineages. Synthesizing sensu lato), is by far the most speciose and the wealth of biological information available phyletically diverse infraorder of Hemiptera in the vast literature on Cicadomorpha and and comprises a substantial proportion viewing this information within a phylogene- (perhaps 6-10%) of the fauna of plant-feeding tic context is no easy task and this paper is by insects. These insects have inhabited Earth for no means an exhaustive review. Given our at least 280 million years (SHCHERBAKOV still rudimentary knowledge of cicadomorp- 1996) and, therefore, have evolved coinciden- han phylogenetic relationships and the diffi- tally with the major lineages of plants, the culty of testing adaptive hypotheses within a development of complex terrestrial ecosy- single lineage (MlTTER et al. 1988), the evolu- stems, the mass extinctions at the - tionary scenarios presented here are largely and -Tertiary boundaries, speculative. Nevertheless, by briefly summari- the break-up of Pangea and countless smaller- zing current knowledge and theories of the scale geologic events that have shaped the evolution of the group, 1 hope to highlight present-day terrestrial realm. Today cicado- gaps in our understanding and topics deser- morphans are ubiquitous in terrestrial habitats ving of further investigation. from equatorial rainforests to the tundra and from sea level to the high mountains, where- Origin of Major Cicadomorp- ver vascular plants can be found. The approxi- han Lineages mately 30,000 described species have been classified into over 5,000 genera and 13 fami- As for most groups, the fossil record lies. Recent sampling in rainforest canopies of Cicadomorpha is extremely fragmentary suggests that the true number of extant cica- (BEKKER-MIGDISOVA 1962, HAMILTON 1992, domorphan species may be ten times higher SHCHERBAKOV 1996); thus reconstructions of (HODKINSON & CASSON 1991, Dietrich unpu- cicadomorphan evolution based on blished). remain highly speculative. Most fossils consist Evidence from various sources may contri- of wing impressions, but the few whole-body bute to an understanding of how cicadomorp- fossils known, scattered across the geologic hans were able to acheive their status as one of time scale from the Permian to the Tertiary, the dominant groups of insect herbivores. provide vital clues regarding the diversity of Phylogenetic studies of modern taxa help elu- morphological forms through time and the cidate the relationships among lineages and timing of the acquisition of certain key inno- their relative ages, and provide a framework vations. Fossil taxa relevant to our understan- for studying the evolution of various adaptati- ding of Cicadomorpha phytogeny continue to ons. Studies of the fossil record provide a be discovered and described, particularly faun- means to correlate the timing of the origin as from the Cretaceous (e.g., HAMILTON 1990, and diversification of various lineages with the 1992) and Tertiary (DIETRICH & VEGA 1995, geological events that shaped our planet, reve- SZWEDO & GEBICKI 1998, SZWEDO & KULICKA al trends in the evolution of the morphologi- 1999), when many modern families and gene- cal traits, and provide information on the ra apparently arose. SHCHERBAKOV (1992, ancient environments in which these insects 1996, and this volume) reviewed the fossil evolved. Studies of the behavior, physiology, evidence pertaining to the evolution of and ecology of modern Cicadomorpha also . provide crucial information relevant to our The fossil record indicates that by the understanding of the adaptations that contri- middle Permian, Cicadomorpha (sensu lato— buted to their evolutionary success and the a paraphyletic assemblage; SHCHERBAKOV means by which closely related species are 1996) were already one of the dominant able to coexist within the same habitats. groups of insect herbivores. These early cica-

156 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at domorphans resembled modern leafhoppers in with an inflated frontoclypeus, presumably having well developed jumping abilities. indicating a shift to feeding on xylem. The Nymphs associated with these fossil taxa were acquisition of enlarged cibarial dilator muscles bizarre biscuitlike creatures that were probab- enabled this group (Clypeata, sensu SHCHER- ly sessile. The apparently explosive diversifi- BAKOV 1996 = Cicadomorpha, sensu stricto) cation of these early hemipterans may have to exploit a new food resource and may have been facilitated by their ability to exploit a facilitated the diversification of this lineage novel food resource due to the acquisition of into what are now recognized as the three piercing-sucking mouthparts, although such modern cicadomorphan superfamilies: Cica-

Fig. 1: acADoroEA CERCOPOIDEA MEMBRACO1DEA Provisional estimate of phylogenetic relationships among the major linea- * Cicadellidae ges of Cicadomorpha based on infor- a mation from DIETRICH & DEITZ (1993), ida e J da e "O idae * rida e \ u I o. C3 o. CM DIETRICH (1999), DIETRICH et al. (2001a, tj o o te r S 8 •o o b). HAMILTON (1999), MOULDS (1999), C3 o. 'D. o. | 5 "ion op •5 a O o JZe I o RAKITOV (1998) and SHCHERBAKOV (1996). yl i la s u ic a u First occurrences of some possible key U 08.

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these analyses indicate that Auchenorrhyncha ly have very loud, persistent calls that facilita- gave rise to Heteroptera + , it is te the attraction of females over greater not clear whether this lineage is a sister group distances. In other groups (e.g., Tettigomyi- of Fulgoromorpha or of Cicadomorpha. ini), the males have shorter, quieter calls and Attempts to reconstruct the phylogeny of the actively seek out the females, some of which major lineages of Cicadomorpha based on are flightless (VlLLET 1999). These alternative morphological data (e.g., HAMILTON 1981, strategies may have evolved in conjunction 1999, EMELJANOV 1987, SHCHERBAKOV 1996, with alternative predator avoidance strategies YOSHIZAWA & SAIGUSA 2001) have yielded and possible tradeoffs between mobility and conflicting results. A more stable and compre- fecundity. Various authors (reviewed by hensive phylogenetic framework, incorpora- MOULDS 1990) have noted that species of ting representatives of relict cicadomorphan cicadas are often associated with particular lineages such as and Tettigarct- habitats or host plants. Thus diversification in idae, and combining morphological and mole- some groups probably resulted from host or cular data, is needed before various alternative habitat shifts. Cicadas are especially diverse in evolutionary scenarios may be tested for Cica- desert environments, where they are often domorpha as a whole. The tree diagram in Fig. active at ambient temperatures that would kill 1 represents a consensus phylogenetic estima- other insects. Recent physiological studies te for Cicadomorpha (sensu stricto) based on (TOOLSON & HADLEY 1987, TOOLSON 1987, my interpretation of the available morpholo- TOOLSON & TOOLSON 1991, SANBORN et al. gical and molecular evidence. 1992) have shown that some cicadas are facul- tatively endothermic, alternatively warming Diversification of Families, themselves through shivering movements of Genera, and Species the flight and/or tymbal muscles and cooling themselves by releasing excess water through Cicadoidea pores on the thorax. This enables these insec- ts to remain active at temperature extremes Based on the fossil record, Cicadoidea that would induce torpor in other insects. apparently arose during the late Triassic or Unfortunately, such physiological adaptations early . Mesozoic cicadas previously pla- have so far been documented in only a few ced in an extinct family (Cicadoprosbolidae) species and, thus, their distribution among are now considered to belong to Tettigarctidae various cicada lineages remains unknown. (BEKKER-MIGDISOVA 1962, SHCHERBAKOV Moreover, phylogenetic studies have so far 1996), a family presently represented by two been performed on only a few groups. Thus, relict southern Australian species (Fig. 2). the extent to which such physiological adap- (sensu lato, Fig. 3) are first recorded tations facilitated cicada diversification needs from the (BEKKER-MlGDISOVA further investigation. 1962). Phylogenetic studies of modern cicada genera and species, ongoing during the past 30 Cladistic biogeographic studies on cicadas, years, have yielded many insights into the fac- particularly in the Indo-Pacific region, indica- tors that influenced the diversification of this te patterns of diversification strongly correla- group. A comprehensive morphology-based ted with geographical vicariance (DUFFELS phylogenetic analysis of the family-group taxa 1986, de BOER 1995, DUFFELS & de BOER is underway and the higher classification will 1996). For example, the main speciation be revised in the near future (MOULDS 1999). events in a cladogram for the tribe Chlorocy- Nevertheless, the restricted distributions of stini correspond with the hypothesized certain currently recognized family-group taxa sequence of fragmentation of the Outer Mela- suggest that continental drift played a role in nesian island arc, and its incorporation into their origin and diversification. Tribes and present-day New Guinea (de BOER 1995). subfamilies also tend to differ from one ano- Speciation within the endemic New Zealand ther in certain behavioral strategies. For Maoricicada appears to have coincided example, in some tribes (e.g., Platypleurini) with the invasion of various montane regions searching during courtship is done primarily following Pleistocene glaciation (BUCKLEY et by females; the males are sedentary and usual- al. 1997).

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The apparently unique life history strategy dators become satiated long before the popu- of the North American periodical cicada lation of cicadas becomes depleted, ensuring genus Magicicada (Fig. 3) deserves special that many cicadas remain to reproduce. Rese- mention. By acquiring an attenuated life- arch is ongoing on the factors that contributed cycle, emerging only every 13 or 17 years, the- to the diversification of Magicicada into the se cicadas mitigate the effects of seven species and 15 allochronic broods cur- through mass emergences that overwhelm the rently recognized (reviewed by SlMON 1988, ability of local predator populations to utilize MARSHALL 2001). Geographic and temporal them as a food resource (KARBAN 1982). Pre- isolation as well as reproductive character dis-

Figs 2-8: Cicadoidea and Cercopoidea: (2) Tettigarcta crinita DISTANT (Tettigarctidae) — cicadas in this family are nearly indistinguishable from Mesozoic fossil taxa; (3) Magicicada Cassini (FISHER) (Cicadidae), a periodical cicada from eastern North America; (4) Paraphilaenus paral- lelus (STEARNS) (Aphrophoridae); (5) obtusa (SAY) (); (6) Pectinahophyes reticulata (SPANGBERG) (Macherotidae); (7) Tomaspis sp. (Cercopidae); (8) spittle mass made by of spumarius (L.) (Aphrophoridae).

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placement have all been proposed as speciati- those genera and species in which fixed nitro- on mechanisms in this genus (ALEXANDER &. gen is transported in the xylem as amino acids MOORE 1962, LLOYD &. DYBAS 1966a, b, or amides. In contrast, fabaceous plant taxa SIMON 1988, MARTIN & SIMON 1988) and with xylem containing ureide nitrogen com- recent population genetic studies have provi- pounds appear to be avoided by these insects. ded support for these mechanisms (MARSHALL Clastopteridae prefer actinorhizal plants (i.e., &. COOLEY 2000, SIMON et al. 2000). those associated with the root-nodule symbi- ont Frankia). Cercopidae exhibit a strong pre- Cercopoidea ference for associative nitrogen fixing grasses (which lack true mycorhizae but grow in loose Of the three major lineages of Cicadomor- association with nitrogen-fixing fungi and pha, Cercopoidea is the least studied phyloge- bacteria). These preferences appear to be con- netically. No comprehensive phylogenetic firmed by the large number of cases of native analysis of the superfamily has ever been spittlebugs that have become pests of introdu- attempted and only a few cladistic analyses of ced or native legumes, or actinorhizal plants genera and tribes have been published (e.g., like Casuarina (THOMPSON 1994). Such corre- LlANG 1998). Thus, the factors that influen- lations between feeding preference and taxo- ced spittlebug diversification remain unclear. nomic affiliation suggest that shifts in feeding Cercopoidea first appear in the fossil record preference may have been involved in the during the Jurassic. Diversification at the divergence of the major lineages of Cerco- family level apparently occurred during the poidea. Such shifts presumably involved Cretaceous and Tertiary (BEKKER-MlGDlSOVA acquisition of a suite of traits including specia- 1962). The four currently recognized families, lized oviposition behaviors and modification Aphrophoridae (Fig. 4), Cercopidae (Fig. 7), of the flora of endosymbionts to accommoda- Clastopteridae (Fig. 5), and te differences in the chemical composition of (Fig. 6), exhibit some differences in behavior the xylem sap of their host plants. A robust and habitat preference. Cercopidae (sensu phylogenetic framework for the superfamily is stricto) occur largely in grasslands and nymphs needed before the roles of such shifts can be of most species apparently feed on grass roots. rigorously tested. Nymphs of the other three families generally As in other cicadomorphans, many cerco- occur on aboveground parts of their hosts. In poid subfamilies, tribes, genera, and species Aphrophoridae, Cercopidae, and Clastopteri- are apparently limited not only in their geo- dae, nymphs live within spittle masses (Fig. graphic ranges (METCALF 1961, 1962) but also 8). In Machaerotidae, nymphs live immersed in their host plants and habitats (HAMILTON in fluid within calcareous tubes cemented to 1982). Thus geographic vicariance and host their host plant. These differences suggest that and habitat shifts presumably played a role in the origins of some major lineages of Cerco- their diversification. In the absence of phylo- poidea involved shifts in habitat and physiology. genetic studies, however, the relative impor- tance of these factors in the origin of spittle- THOMPSON (1994) summarized the still bug lineages remains unknown. rather sparse knowledge of host plant and eco- logical associations among Cercopoidea, noting that at least three of the four families, Membracoidea although having broad host ranges overall, Comprising approximately 25,000 descri- exhibit statistically significant preferences for bed species placed in 3,500 genera, 150 tribes, particular groups of plants. With the possible 50 subfamilies, and 5 extant families (DEITZ & exception of Machaerotidae, spittlebugs exhi- DIETRICH 1993, HAMILTON 1999), Membra- bit a strong preference for nitrogen-fixing coidea is by far the largest and most phyleti- plants. Interestingly however, different cerco- cally diverse of the extant cicadomorphan poid families apparently prefer different groups superfamilies (Figs 10-21). The ecology and bio- of such plants. Aphrophoridae exhibit a nomics of Membracoidea have been studied strong preference for legumes, particularly intensively due to their considerable econo-

160 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at mic importance, but explicit phylogenetic membracoids also produced brochosomes estimates based on cladistic analysis are avai- (see Rakitov, this volume), but the absence of lable for only a few groups (OMAN et al. 1990). brochosomes in Myerslopiidae, the most The earliest known membracoids (Karajassi- plesiomorphic extant family, suggests that dae from the lower Jurassic) had inflated faces, brochosomes were acquired in ancestral Cica- indicating that they fed on xylem, like their dellidae (Fig. 1). Modern leafhoppers anoint cicada and spittlebug relatives (SHCHERBAKOV themselves with brochosomes after molts and 1992). These membracoids had, however, the brochosome coating provides an extremely already acquired marked modifications to hydrophobic coating, protecting the insects

Mi««Ms»

Egg guarding I I absent i^H present

their leg and forewing structure, implying from becoming trapped in water droplets and Fig. 9: Cladogram of Membracidae from the morphology-based phyloge- their own copious excreta (RAKITOV 1996, behavioral shifts that further distinguished netic analysis of DIETRICH et al. (2001b) them from other cicadomorphans. The for- 1998, 1999, and this volume). showing the distribution of egg-guar- ewing became narrower, particularly in the The modifications of the legs of adults ding behavior. Egg guarding is plesio- morphic in treehoppers, occurring in costal area, and the costal margin no longer and nymphs and production of brochosomes Aetalionidae and the plesiomorphic extended ventrad of the thoracic pleura. The provided membracoids with much greater membracid subfamily Endoiastinae. This behavior was apparently lost ear- hind coxae became enlarged and the jumping mobility and apparently gave them access to ly in the evolution of Membracidae, muscles became more highly developed. The microhabitats not available to their cicada but was later regained in various hind femora elongated and the tibiae acquired and spittlebug relatives. For example, the abi- lineages. Ant mutualism (not mapped) is much more widespread among longitudinal rows of enlarged spinelike setae. lity to live exposed on aboveground plant membracids, but occurs sporadically in The acquisition of specialized leg chatotaxy parts may have helped facilitate the shift from most groups that lack egg guarding implies that early membracoids „anointed" the plesiomorphic xylem-feeding strategy behavior. Ant-mutualism has been lost independently in Aconophorini and (retained by cicadelline leafhoppers) to the themselves with products of the Malpighian Hoplophorionini, the two membracid tubules, using the rows of setae to spread the phloem-feeding strategy that predominates in tribes exhibiting the most highly deve- secretion over the integument. Possibly, early the modern membracoid fauna, as well as the loped parental care behavior.

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shift to parenchyma feeding in Typhlocybinae. rence of ant-mutualism and parental care Membracoid nymphs feeding exposed on lea- among aetalionids and membracids suggests ves and twigs faced a new set of challenges, that acquisition of these behaviors coincided including increased exposure to predators and with the origin of the treehopper lineage that more extreme fluctuations in environmental now comprises the vast majority of species. (particularly microhabitat) conditions. Diffe- Phylogenetic analysis of Membracidae indica- rent lineages have met these challenges in tes that ant-mutualistic behavior became spo- myriad ways. To avoid predation, some rely on radic or, more rarely, was completely lost in or (reviewed by Mejda- various treehopper lineages (e.g., Stegaspidi- lani et al., this volume), some on agility, nae, Nicomiinae, Darninae, Ceresini, Smili- others on defensive morphological armature ini). In some, e.g., and Dar- (EKKENS 1972), and still others on mutualistic ninae, the nymphs became solitary and acqui- associations with aggressive social hymenop- red crypsis-enhancing morphological features terans (WOOD 1984). In some groups, particu- (Dietrich et al. 2001b). In others, e.g., Aco- larly Membracidae, different strategies are nophorini (Fig. 18) and Hophlophorionini, employed by adults and immatures; thus selec- complex parental care behaviors provided tion led to greater divergence between the alternative means for protecting nymphal adult and nymphal body forms (e.g., Fig. 20). aggregations (DIETRICH & DEITZ 1991; MCKA- Cladistic analyses of the major membraco- MEY & DEITZ 1996). id lineages indicate that some life history traits Acquisition of such alternative defensive are phylogenetically conservative, suggesting strategies may account for at least some of the that their acquisition coincided with the ori- major divergences within Membracoidea but gins of major lineages. For example, with the they alone do not explain why there are so exception of Eurymelinae (sensu stricto, many species of leafhoppers and treehoppers. EVANS 1931), ant-mutualism appears to be Data on host associations, geographic range, rare among leafhoppers, occurring sporadical- and habitats, coupled with phylogenetic ana- ly within a few other subfamilies (e.g., Idioce- lyses at the genus and species level within rinae, Macropsinae, ; DlE- several groups suggest numerous scenarios that TRICH & McKAMEY 1990). In contrast, ant- might account for recent divergences within mutualism is common in the treehopper fami- membracoid genera. Estimates of the phylo- lies Aetalionidae (Fig. 15) and Membracidae, geny of Cicadellidae based on morphology and is often associated with parental care (egg (DIETRICH 1999) and DNA sequences (DIE- guarding, Figs 14, 18) and gregarious behavior TRICH et al. 2001a) indicate that one of the (WOOD 1984). Among treehoppers, ant- major lineages of leafhoppers (comprising Del- mutualism and parental care are plesiomor- tocephalinae and several other subfamilies) phic traits (Fig. 9), occurring nearly universal- comprises species largely associated with ly in Aetalionidae and in the most plesiomor- semiarid or arid habitats (particularly gras- phic membracid subfamily Endoiastinae (Die- slands). Thus, a habitat shift from forests to trich et al. 2001b). Most other membracid grasslands may have coincided with the origin subfamilies (Stegaspidinae, Centrotinae, of this lineage. Other major membracoid Heteronotinae, , Darninae, and lineages exhibit a wide variety of host and ) contain at least some ant-mutuali- habitat preferences, suggesting more complex stic species and ant-mutualism appears to patterns of evolutionary diversification. occur universally within some tribes, e.g., Amastrini and Tragopini. The behavior of the Because host and habitat associations are most plesiomorphic treehoppers, Melizoderi- often fairly conservative within genera of dae (sister group of Aetalionidae + Membraci- Membracoidea, biogeographic vicariance has dae; endemic to ; Fig. 13), has not been often been invoked to explain recent speciati- studied, but the morphology of melizoderid on events. In Enhomus, a genus of large, nymphs indicates that they are adapted for flight-limited leafhoppers endemic to north- crypsis and may not be ant-mutualistic (DlE- western North America, partitioning of the TRICH & DEITZ 1993). The widespread occur- range of the host plant (Balsamorrhyza)

162 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at appears to have resulted, in part, from vicari- Studies of several groups of grass- and sed- ant processes associated with glaciation and ge-feeding deltocephaline leafhoppers indica- river capture in the Columbia River basin te that host plant shifts were involved in the (OMAN 1987, HAMILTON & ZACK 1999). Phy- diversification of individual genera and spe- logeographic studies incorporating mitochon- cies. DIETRICH et al. (1997) found that several drial DNA sequence data are needed to test of Flexamia species (Fig. 12) were asso- such hypotheses which, thus far, have been ciated with particular grass genera or species. based on morphological and geological data Similar patterns appear in Athysanelia, a large alone. genus of mostly flightless leafhoppers (HlCKS

Figs 10-15: Membracoidea: (10) Myerslopia chilensis NIELSON (Myerslopiidae) — myerslopiids, restricted to Chile and New Zealand, are thought to be a sister group to the lineage comprising the remaining families of Membracoidea; (11) Hylaius oregonensis (BAKER) (Cicadellidae: Err- homeninae); (12) Flexamia grammica (BALL) (Cicadellidae: Deltocephalinae), a grass-specialist ; (13) Llanquihuea pilosa LIN- NAVUORI & DELONG (Melizoderidae) — melizoderids, restricted to Chile, are thought to be the sister group of the lineage comprising the remaining two treehopper families Aetalionidae and Membracidae; (14) a female Aetalion reticulatum (L) (Aetalionidae: Aetalioninae) guarding an egg mass; (15) ant-attended aggregation of Tropidaspis sp. (Aetalionidae: Biturritiinae).

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et al. 1988, WHITCOMB et al. 1986). Plesio- ra species may occur together on the same morphic species of Daündus specialize on Trip- plant. ROSS (1957b, 1958) suggested that sacum spp., but shifts to feeding on Zea spp. many such species have identical niches, an occurred independently in two lineages (TRIP- apparent contradiction of Gause's Law. But LEHORN & NAULT 1985, DIETRICH et al. 1998). MCCLURE & PRICE (1975, 1976) demonstra- Within such host-associated lineages, closely ted that, at least in sympatric sycamore-fee- related species tend to be allo- or parapatric, ding species, competition can be suggesting that host shifts followed by biogeo- severe. Their experiments indicated that a graphic partitioning of the host range could combination of biogeographic and microhabi- account for much of the diversification within tat partitioning may reduce interspecific com- the genus. Diversification of Limotettix (sensu petition and facilitate coexistence among the lato) may have involved a combination of species whose ranges overlap. Other groups of host and habitat shifts (e.g., among various closely related, coexisting leafhopper and tre- kinds of wetlands and wetland plants; HAMIL- ehopper species (e.g., the oak-feeding Cyrtolo- TON 1994). bus generic group of North American treehop- In the above mentioned genera, host plant pers), appear to show similar patterns. Phylo- shifts occurred mostly among closely related genetic studies of some such groups are under- plant species (e.g., grasses). In other membra- way (e.g., WOOD et al., pers. comm.). coid genera, different species specialize on Character displacement has also been unrelated plants that differ substantially in invoked to explain the coexistence of closely their phenologies. WOOD'S (1992 and refs.) related species in the same habitat (e.g., extensive studies of one such group, the HAMILTON 1998, 2000). Studies of acoustic binotata species complex of North courtship signals (e.g., HEADY et al. 1986, American treehoppers, have revealed patterns HUNT 1994, TISHECHKIN 1998, 2000a, b) and of diversification consistent with a sympatric the extensive divergence in male genitalia, speciation model. Isolation of new host-asso- particularly in leafhoppers, appear to confirm ciated Enchenopa lineages may have been the role of sexual selection in reinforcing bar- mediated by the phenological differences riers to interbreeding among congeners. among their various host plants, resulting in Indeed, within most membracoid genera, asynchronous development and temporal iso- other aspects of the morphology are highly lation of populations within a small geogra- conservative, although multivariate morpho- phic area. In the E. binotata complex such a metric analysis of „external" structures (e.g., process has apparently given rise to at least WOOD & PESEK 1992, DIETRICH & POOLEY seven biological species. Similar mechanisms 1994) may be capable of distinguishing species may explain diversification within some linea- in some groups. Divergence associated with ges of typhlocybine and deltocephaline leaf- the most recent speciation events in Membra- hoppers, where numerous cryptic, host-specia- coidea is often manifested in subtle physiolo- list species have been discovered among speci- gical, ecological, and behavioral differences, mens of what were once thought to be single in the absence of substantial modification of polyphagous species (e.g., DELONG 1931, the morphology, and many cryptic species Ross 1957a, 1965, HEPNER 1966, KLEIN & undoubtedly await discovery. GAFNY 1999).

Although a combination of host and habi- The Membracid Pronotum: An tat shifts may suffice to explain the diversifi- Evolutionary Enigma cation of many membracoid genera and spe- cies, the present geographic ranges of many One aspect of cicadomorphan evolution closely related species overlap substantially that continues to defy explanation is the and it is not uncommon to find congeneric extreme divergence in pronotal shape obser- species occupying the same habitats feeding ved among genera of Membracidae (Figs 16- on the same host plant. For example, HEPNER 21). Early observers (e.g., POULTON 1891, (1976) noted that as many as 100 Erythroneu- MANN 1912) suggested that these insects

164 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at mimic various parts of their host plants al projections of many species deter venebrate (thorns, seeds, bark, etc.) or venomous insects predators, and this hypothesis has received (, ). HAVILAND (1926), in addition some support by observations of later workers to providing evidence of crypsis and mimicry (e.g., EKKENS 1972; WOOD 1975). FUNKHOUSER in some species, suggested that some strikingly (1951), while noting the possibile adaptive patterned species are aposematic. WOOD significance of some pronotal shapes, sugge- (1975) confirmed that two North American sted that the pronotum of many species lacked species exhibit aposematic coloration. adaptive significance and attributed the diver- Haviland also suggested that the spiny pronot- sity of bizarre forms to orthogenesis (i.e., evo-

Figs 16-21: Membracidae: (16) a member of the Enchenopa binotata (SAY) species complex of the eastern U.S.A.; (17) Cladonota sp., Guyana; (18) female Aconophora sp. guarding eggs and nymphs, Peru; the bands of sticky secretion coating the stem on either side of the egg mass apparently deter predators and parasitoids—note the trapped on the left; (19) globulare (F.), Guyana; (20) Heteronotus sp. from Ecuador; the strikingly marked adult of this species contrasts sharply with the cryptic nymph (exuviae at lower right); (21) extrema BALL, eastern U.S.A.

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lution along a progressive line independent of Literature Cited natural selection). STRÜMPEL (1972) refined this idea by demonstrating that differences in ALEXANDER R. S T. MOORE (1962): The evolutionary relationships of 13- and 17-year periodical cica- pronotal shape among species in some genera das, and 3 new species. — Mus. Zool. Misc. Publ. may be explained by simple allometric growth 121, Univ. Michigan, Ann Arbor, USA. models. More recently, WOOD & MORRIS BEKKER-MIGOISOVA E.E. (1962): Superorder Rhynchota. (1974) noting the presence of numerous sen- Insects with proboscis. In Rohdendorf, B.B. Fun- damentals of Paleontology, vol. 9, Arthropoda, sory pits on the membracid pronotum and sug- Tracheata, Chelicerata. Akademiya Nauk SSSR, gested that the expansion of the pronotum Moscow [In Russian; English translation 1991, Smithsonian Institution, Washington, DC, USA.) may have been an adaptive response to selec- tion for increased sensory surfaces or evapora- BOER A.J. de (1995): The phylogeny and taxonomic status of the Chlorocystini (sensu stricto) tive surfaces for dispersal of pheromones. (Homoptera, Tibicinidae). — Contr. Zool. 64: Another possible function of the expan- 201-231. ded pronotum is suggested by recent physiolo- BUCKLEY T, ARENSBURGER P., CHAMBERS G. & C. SIMON (1997): Testing hypotheses of New Zealand cica- gical studies of thermoregulation in cicadas da evolution: I. Australian relationships and (see Cicadoidea, above). Some cicada species, evolution of Maoricicada. Program and Abstract when subjected to high ambient temperatures, Book, 9th International Auchenorrhyncha Con- gress, 17-21 February 1997. cool themselves by releasing excess water CAMPBELL B.C., STEFFEN-CAMPBELL J.D., SORENSEN H.T & through pores on the thorax and abdomen. If R.J. GILL (1995): Paraphyly of Homoptera and similar cooling mechanisms are present in the Auchenorrhyncha inferred from 18S rDNA related Membracidae, then selection may nucleotide sequences. — Syst. Entomol. 20: 175- 194. have favored pronota with larger, pitted surfa- DEITZ L.L. & C.H. DIETRICH (1993): Superfamily Mem- ce areas to provide greater efficiency in heat bracoidea (Homoptera: Auchenorrhyncha). I. transfer. This possible physiological function Introduction and revised classification with new of the enlarged membracid pronotum merits family-group taxa.—Syst. Entomol. 18: 287-296. investigation. Membracidae achieve their DELONG D.M. (1931): A revision of the American spe- cies of Empoasca known to occur north of Mexi- greatest diversity in tropical rainforests, savan- co. — U. S. Dept. Agr. Tech. Bull. 231: 1-59. nas, and deserts, environments where, given DIETRICH C.H. (1999): The role of grasslands in the their tendency to be active during the day, diversification of leafhoppers (Homoptera: Cica- they may be subjected to extreme heat. Impro- dellidae): A phylogenetic perspective. Proc. 15th North American Prairie Conf., Oct 23-26, 1996, ved thermoregulatory efficiency provided by St Charles, IL, USA, pp. 44-49. the enlarged pronotum may have facilitated DIETRICH C.H. & L.L. DEITZ (1991): Revision of the diversification in treehoppers by giving them a Neotropical Treehopper Tribe Aconophorini selective advantage in such environments, (Homoptera: Membracidae). — North Carolina enabling them to engage in courtship activity Agr. Res. Serv. Tech. Bull. 293: 1-134. during midday when many of their predators DIETRICH C.H. & L.L. DEITZ (1993): Superfamily Mem- bracoidea (Homoptera: Auchenorrhyncha). II. are inactive. Cladistic analysis and conclusions. — Syst. Entomol. 18: 297-312. DIETRICH C.H. & S.H. MCKAMEY (1990): Three new idio- Acknowledgments cerine leafhoppers (Homoptera: Cicadellidae) from Guyana with notes on ant-mutualism and subsociality. — Proc. Entomol. Soc. Washington 1 am indebted to L.L. Deitz, K.G.A. 92: 214-223. Hamilton, S.H. McKamey, R.A. Rakitov, D.E. DIETRICH C.H. S CD. POOLEY (1994): Automated iden- Shcherbakov, M.D. Webb, R.F. Whitcomb, tification of leafhoppers (Homoptera: Cicadelli- T.K. Wood, and other colleagues too nume- dae: Draeculacephala). — Ann. Entomol. Soc. Amer. 87:412-423. rous to mention for many helpful discussions DIETRICH C.H. & F.E. VEGA (1995): Leafhoppers that have helped shape my views of cicado- (Homoptera: Cicadellidae) from Dominican morphan evolution. R.A. Rakitov and D.M. amber. — Ann. Entomol. Soc. Am. 88: 263-270.

Takiya critically reviewed the manuscript. DIETRICH C.H., WHITCOMB R.F. & W.C. IV. BLACK (1997): This work was supported in part by grants Phylogeny of the North American grassland leafhopper genus Flexamia (Homoptera: Cica- from the National Science Foundation (DEB- dellidae) based on mitochondrial DNA sequen- 9726282, 9978026, and 0089671). ces. — Mol. Phylogenet. Evol. 8: 139-149.

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Address of the author:

Dr. Christopher H. DIETRICH Center for Biodiversity, Illinois Natural History Survey 607 E. Peabody Dr., Champaign, Illinois 61820, USA [email protected].

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