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An Overview of Fulgoromorpha and Cicadomorpha in East African Copal (Hemiptera) 57-66 © Biologiezentrum Linz/Austria; Download Unter 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): Stroinski Adam, Szwedo Jacek Artikel/Article: An overview of Fulgoromorpha and Cicadomorpha in East African copal (Hemiptera) 57-66 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at An overview of Fulgoromorpha and Cicadomorpha in East African copal (Hemiptera) A. STROINSKI & J. SZWEDO Abstract East African copal is rich in inclusi- ons, which normally do not occur single, but the number of species described from East African copal is relatively low. Inclu- sions of Fulgoromorpha and Cicadomor- pha (Hemiptera) in East African copal are not very frequently mentioned. The age of this fossil resin - copal - is still uncertain, it may come from the Pliocene, but is generally regarded as Pleistocene (1-5 Ma). Key words: Hemiptera, Fulgoromor- pha, Cicadomorpha, Ricaniidae, Derbidae, Issidae, Cicadidae, copal, East Africa, Pleistocene. Denisia 04, zugleich Kataloge des OÖ. Landesmuseums, Neue Folge Nr. 176 (2002). 57-66 57 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Introduction resins, or to all fossil resins except Baltic amber. 'Copal' is sometimes used in place of Planthoppers - Fulgoromorpha, and Cica- 'resin', both for recent 'copal' and for fossil or domorpha - treehoppers, frog-hoppers, cicadas subfossil 'copal', i.e. older than Pleistocene and leafhoppers - are hemipterans common in (SCHLÜTER &. von GNIELINSKI 1987). Accor- almost all habitats throughout the world. It is ding to SCHLEE (1984, 1990), the term 'copal' estimated that more than 43.000 species of is also used for resins younger than 250 years! Fulgoromorpha and Cicadomorpha have been On the other hand, purists reserve this name described (OMAN & SAILER 1986). for the resin of Caesalpiniaceae (GEIRNAERT The fossil record of the representatives of 1998). Regarding the stratigraphic position of these hemipteran lineages reaches the end of resins called 'copals', it seems that - in spite of the Early Permian (SHCHERBAKOV 1996, all these nomenclature uncertainties and the 2000). The oldest resin inclusions of these ins- lack of a strict definition of copal - the term ects come from Lower Cretaceous Lebanon refers to subfossil resins originating from the amber (FENNAH 1987). Two species of plant- Southern Hemisphere and of Pleistocene age hoppers are known from Late Cretaceous Bur- (1.64-5.2 Ma). mese amber (COCKERELL 1917). Representati- Where are these subfossil resins located? ves of both Fulgoromorpha and Cicadomor- Most subfossil resins occur in tropics or very pha lineages are quite common in Eocene Bal- wet temperate areas, where the trees produ- tic amber inclusions (SZWEDO & KULICKA cing resin are still present. A single geographi- 1999). The first description of representatives cal region may have copal deposits of various of this family from Baltic amber comes from ages, ranging from recent to many thousands the second half of the 19th century (GERMAR years old. & BERENDT 1856), later followed by BERVOETS (1910), COCKERELL (1910), JACOBI (1938), On North Island of New Zealand the resin USINGER (1939), EMELJANOV (1990, 1994), was and continues to be exuded from the GEBICKI & SZWEDO (2000a, b), and SZWEDO & Araucariaceae species Agathis australis - the GEBICKI (1998, 1999, 2000). In a piece of Bit- Kauri pine. This resin is also called 'Kauri terfeld amber from Germany, dated Miocene gum'. The stratigraphic age of this resin is but recently regarded as redeposited and belie- obscure (SCHLÜTER & VON GNIELINSKI 1987). ved to be of Eocene age (WEITSCHAT 1997), a During the Mesozoic, Araucariaceae were few inclusions were found, but not yet formal- common throughout.the Northern Hemisphe- ly described. From Oligocene/Miocene New re, but disappeared in the Early Tertiary, so the World resins a few representatives planthop- recent distribution is now restricted to the pers and leafhoppers have been mentioned Southern Hemisphere. The age of Kauri copal (FENNAH 1963, GEBICKI & WEGIEREK 1993, ranges from at least the Eocene up to the pre- DIETRICH & VEGA 1995, SZWEDO 2000a, sent (THOMAS 1968, SCHLÜTER 1978). This STROINSKI & SZWEDO 2000, EMELJANOV & kind of resin is also known from Australia SHCHERBAKOV 2000, SZWEDO & STROINSKI (HILLS 1957) and the Philippines (DURHAM 2001). 1956). Fossil resin has also been reported from 1. Copal - fossil resin and its signi- Indonesia (DURHAM 1956), probably origina- ficance ting from a tree of the family Dipterocarpa- Fossil, subfossil and recent resins, inclu- ceae, and dated Upper Miocene - Sumatra ding these known as copal and amber, are localities, or Pliocene - Java localities widespread throughout the world. The word (SCHLÜTER & VON GNIELINSKI 1987). More- 'copal' is derived from Mexican Spanish, from over, fossil resins with inclusions have been Nahuatl word 'copalli' (RICE 1987, AHD reported from Celebes (KLEINE 1924, who 2000). There is some confusion concerning described fossil Coleoptera: Scolytidae) and the term, because it is sometimes used to refer from the Lower-Mid Miocene of the Sarawak to any resin formed by tropical legume trees Province in Malesia (HlLLMER, VOIGT & and the araucarians, as well to any subfossil WEITSCHAT 1992). 58 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at So called "Mizunami amber" is recorded genera, and probably also other Caesalpinia- from Japan. It was supposed to date back to ceae, originated during the Early Cretaceous the Middle Pleistocene, but according to on the joined South American-African land- radiometric data it is much younger, only mass. After the separation of the continents, about 33 000 (-1600 to +2000) years old they were subject to individual evolutionary (SCHLEE 1984). processes based on genetic changes and envi- From Israel another fossil resin has been ronmental selection (PoiNAR 1992). recorded. It is probably of Pleistocene age and The East Coast of Africa (southern parts originates from Pistatia lenasculus tree, of the of Kenya and Tanzania) is one of the seven family Anacardiaceae (SCHLÜTER & VON plant species endemism and richness areas in GNIELINSKI 1987). sub-Saharan Africa (LlNDER 2001). An analy- Most copals derive from leguminous sis of the inclusions of living organisms found (Fabales) trees of the family Caesalpiniaceae, in fossil and subfossil resins originating from particularly of the genus Hymenaea. Copal the trees of this area is very important. Com- occurs in Central and South America: Minais parison between recent fauna and fossil orga- Gerais Province in Brazil, Mariquita Province nisms could help in description of life-histo- in Colombia and eastern Dominican Repu- ries of the insects as well as other animals and blic. It is sold to the collectors as "Pliocene plants. Detailed analysis of the characters amber", about 2 Ma old, but in fact it is mere- found in fossil representatives of the groups ly several hundreds years old (GRIMALDI will advance the search of the origins and 1996). polarization of characters used in cladistic A tree of the genus Hymenaea is also pre- analyses. sent in Africa, it is Hymenaea verrucosa regar- ded as a member of the primitive section 2. Inclusions in East African copal Trachybbium, and restricted to East Africa and adjacent islands (PoiNAR 1992). This tree grows Most descriptions of inclusions in fossil in lowland evergreen and semi-deciduous resins come from Eocene Baltic amber and forests. HUEBER & LANGEHEIM (1986) suggest Oligocene/Miocene Dominican and Mexican that the section Trachybbium is of African ori- amber. The specimens stored in the Natural gin, and that Hymenaea verrucosa is a relict of History Museum London, labeled 'amber' and the former Pan-African distribution. shown as amber in the Quarterly Journal of Other African resins have been recorded Science in 1868 are actually East African from West Africa: Guinea, Sierra Leone, Gha- copal (ROSS 1999). Also the American Muse- na, Benin, Gabon, Congo and North East um of Natural History has an extensive collec- Angola (SCHLÜTER & VON GNIELINSKI 1987, tion of insects preserved in East African copal RlCE 1987). These resins originate from an- (GRIMALDI 1996). Also copal from Madagas- other Caesalpiniaceae (Fabales) tree of the car is rich in inclusions (Geinaert, personal genus Copaifera, with the estimated age ran- communication). ging from the Pleistocene to 8000 years B.C. Generally, copal inclusions have rarely (SCHLÜTER & VON GNIELINSKI 1987). It is been described, although East African copal interesting, that only four species of Copaifera seems to be the source of relatively high num- occur in West Africa, and most of them in ber of taxa described (SCHLÜTER & VON South America, particularly in Brazil (LANGE- GNIELINSKI 1987). HEIM 1973). Among the groups mentioned from East Copal from Hymenaea verrucosa - 'Mand- African copal are: Araneae, Scorpionida, rorofo' in the Malgaskan language - has also Blattodea, Isoptera, Embioptera, Psocoptera, been reported from Madagascar (GEIRNAERT Hemiptera, Coleoptera, Hymenoptera, Lepid- 1998). optera, Diptera and Reptilia. A sole species of The present existence and distribution of Hemiptera: Cicadomorpha - Cicada forsythi related resin-producing trees of the genera BUCKT. has been described by Buckton Hymenaea and Copaifera suggest that these (BUCKTON 1890, 1891). 59 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at
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