ZOBODAT - www.zobodat.at Zoologisch-Botanische Datenbank/Zoological-Botanical Database Digitale Literatur/Digital Literature Zeitschrift/Journal: Denisia Jahr/Year: 2002 Band/Volume: 0004 Autor(en)/Author(s): Dietrich Christian O., Dietrich Christian O. Artikel/Article: Evolution of Cicadomorpha (Insecta, Hemiptera) 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 family- poidea and Membracoidea) are one of the group taxa may also have coincided with dominant groups of plant-feeding insects, 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 species, 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 fossil 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: cicada, spittlebug, leafhop- hind legs, specialized grooming structu- per, treehopper, 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), 155-170 155 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at 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- leafhoppers and treehoppers (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 Permian- tionary scenarios presented here are largely Triassic and Cretaceous-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 insect 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 fossils 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 Auchenorrhyncha. 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- \ ges of Cicadomorpha based on infor- J * e Cicadellidae * e a e mation from DIETRICH & DEITZ (1993), ida da "O idae rida u I o. C3 r o. CM DIETRICH (1999), DIETRICH et al. (2001a, tj o o te S 8 •o o b). HAMILTON (1999), MOULDS (1999), C3 o. 'D. o. | 5 "ion •5 O i op a o s a JZe I o RAKITOV (1998) and SHCHERBAKOV (1996). yl la u ic u First occurrences of some possible key U 08. <a.^ U U innovations are indicated (see text for discussion). Extinct taxa are indicated u by an asterisk. Lineages 1-3 of Cicadel- egg-guarding ant-mutualism lidae are defined as follows: Lineage 1 includes Cicadellinae (in part), Makilin- ,—j-, brochosomes giinae, Phereurhininae, Tinterominae, __J lost i nymphs with m i nymphs in Signoretiinae, Phlogisinae, Nirvaninae, fossorial front legs | spittle Coelidiinae, Typhlocybinae; Lineage 2—Euacanthellinae, Aphrodinae, i brochosomes Xestocephalinae, Acostemminae, Neobalinae, Stegelytrinae, Mukariinae, Penthimiinae, Selenocephalinae, Del- tocephalinae, Koebeliinae, Arrugadi- anointing behavior (?) nae, Drakensbergeninae; Lineage 3— Hylicinae, Austroagalloidinae, Ulo- pinae, Agalliinae, Megophthalminae, Adelungiinae, Evansiolinae, Macrop- sinae, Tartessinae, Nioniinae, Idioce- rinae, Eurymelinae, Ledrinae, lassinae, Scarinae. Lineage 3 is paraphyletic mouthparts are also known in some Carboni- doidea (cicadas), Cercopoidea (spittlebugs), with respect to the clade Melizoderi- ferous-age Paleodictyoptera (LABANDEIRA & and Membracoidea (leafhoppers and treehop- dae+Aetalionidae+Membracidae. Err- homeninae (placed as a tribe of Cica- PHILLIPS 1996). Early radiations in Hemiptera pers). dellinae by OMAN et al. (1990)) is of apparently involved modification of the wing Several studies have attempted to assess uncertain position. Tettigarctidae may and leg morphology to provide improved