New Termitaphididae and Aradidae (Hemiptera) in Mexican and Dominican Amber
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Palaeodiversity 4: 51–62; Stuttgart 30 December 2011. 51 New Termitaphididae and Aradidae (Hemiptera) in Mexican and Dominican amber GEORGE POINAR, JR. & ERNST HEISS Abstract A new species of termite bug, Termitaradus dominicanus POINAR, n. sp. (Hemiptera: Termitaphididae) and a new genus and species of fl at bug, Brevisensoria incrustata POINAR, n. gen., n. sp. (Hemiptera: Aradidae) are described from Dominican amber. Termitaradus dominicanus POINAR, n. sp. is distinguished from previously described mem- bers of the genus by lobules with terminal fl abella composed of two to four minute, non-serrate setae, with each setal group embedded in a hardened envelope or deposit. Brevisensoria incrustata POINAR, n. gen., n. sp. can be separated from other members of the Aradidae by the absence of compound eyes, ocelli and wings, modifi ed head fused to the prothorax, unique antennal structure and the dorsum forming a continuous body covering with a peripheral margin bearing lobes. The oval body shape, convex dorsum with incrustations, convex ventrum, absence of lobules and unique structure of the antennae distinguish the new genus from members of the Termitaphididae. In addition, fi ve specimens of Termitaradus protera POINAR & DOYEN in a single piece of Mexican amber that also contains seven worker termites are characterized. K e y w o r d s : Dominican amber, Tertiary termite bugs, Tertiary fl at bug, Mexican amber. Zusammenfassung Aus Dominikanischem Bernstein werden Termitaradus dominicanus POINAR, n. sp. (Hemiptera: Heteroptera: Termitaphididae) und die neue Gattung und Art Brevisensoria incrustata POINAR, n. gen., n. sp. (Hemiptera: Hete- roptera: Aradidae) beschrieben. Termitaradus dominicanus POINAR, n. sp. unterscheidet sich von den anderen Arten der Gattung durch die terminalen Fortsätze (fl abellae) der Lateralloben der Segmente, welche mit zwei bis vier kleinen, nicht gezähnten Borsten besetzt sind und jedes Borstenbündel einer verhärteten Basis entspringt. Bre- visensoria incrustata POINAR, n. gen., n. sp. kann von anderen Taxa der Familie Aradidae durch das Fehlen von Komplexaugen, Ocellen und Flügeln, dem modifi ziertem Kopf, der mit dem Prothorax verwachsen ist, die einzig- artige Fühlerstruktur und die Dorsalseite des Körpers unterschieden werden, deren Segmente mittig durchgehend verschmolzen sind, jedoch laterale Loben ausgebildet haben. Die ovale Körperform, die konvexe Dorsalseite mit Krustenstrukturen und die konvexe Unterseite, das Fehlen von terminalen Fortsätzen der Lateralloben und die ein- zigartige Struktur der Fühler unterscheiden die neue Gattung auch von den Termitaphididae. Weiters werden fünf Exemplare von Termitaradus protera POINAR & DOYEN beschrieben, welche sich zusammen mit sieben Arbeitern einer Termite in einem Stück Mexikanischem Bernstein befi nden. Contents 1. Introduction ........................................................................................................................................................... 51 2. Materials and methods ..........................................................................................................................................52 3. Systematic palaeontology......................................................................................................................................53 Order Hemiptera ...................................................................................................................................................53 Family Termitaphididae ...............................................................................................................................53 Genus Termitaradus MEYERS ...........................................................................................................53 Family Aradidae ...........................................................................................................................................57 Genus Brevisensoria POINAR, n. gen. ..............................................................................................57 4. References .............................................................................................................................................................62 1. Introduction amber indicate an ancient origin for this group, certainly extending back at least to the Cretaceous and probably Termite bugs of the family Termitaphididae (Hemip- even earlier (POINAR & DOYEN 1992). tera) represent extremely modifi ed hemipterans that occur Flat bugs of the family Aradidae have oval to ellip- in termite colonies worldwide. They have lost their eyes soidal dorsoventrally fl attened bodies with porrect heads and wings and the head, thorax and abdomen are fused into bearing prominent antennae. They are found under bark or a protective shield covering the shortened legs and anten- associated with dead wood, presumably obtaining nour- nae. Such structural modifi cations allow them to survive ishment from wood-rotting fungi. Many species have lost in the confi nes of termite nests. The world-wide distribu- their wings and have a rough surface that blends with the tion of extant forms and fossils in Mexican and Dominican surroundings, thus depending on camoufl age rather than 52 PALAEODIVERSITY 4, 2011 fl ight to avoid detection. Some 475 species are known synopsis of Mexican amber are presented in POINAR (1992). from the Neotropics (KORMILEV & FROESCHNER 1987). Amber from this region was produced by Hymenaea mex- The present paper describes a new species of Termit- icana (Fabaceae) (POINAR & BROWN 2002) and occurs in aradus and a new genus and species of aradid, both from lignitic beds among sequences of primarily marine cal- Dominican amber, and briefl y characterizes fi ve new spec- careous sandstones and silt. The amber is associated with imens of Termitaradus protera POINAR & DOYEN, 1992 Balumtun Sandstone of the Early Miocene and the La together with seven worker termites, in Mexican amber Quinta Formation of the Late Oligocene with radiomet- (Fig. 1). ric ages from 22.5 to 26 million years (BERGGREN & VAN C OUVERING 1974). Since the amber is secondarily depos- Acknowledgements ited in these marine formations, it is undoubtedly some- We thank W. WEITSCHAT for the loan of the amber piece with what older than the above dates. The specimen is in the the Mexican termitaradids and ART BOUCOT and ROBERTA POINAR collection of W. WEITSCHAT Hamburg, Germany. for comments on earlier drafts of this manuscript. The amber containing the Dominican specimens origi- nated from mines in the northern mountain range (Cordil- lera Septentrional) of the Dominican Republic, between 2. Materials and methods the cities of Puerto Plata and Santiago. Amber from this deposit was produced by Hymenaea protera POINAR, 1991 The Mexican amber piece is elliptical in shape, meas- (Fabaceae). Dating of Dominican amber is controversial, uring 38 mm in length, 15 mm in width and 12 mm in with the youngest proposed age of 20–15 mya based on depth. The amber originated from the Simojovel area of foraminifera (ITURRALDE-VINENT & MACPHEE 1996) and Chiapas, Mexico. Locations of the Chiapas mines and a the oldest as 45–30 mya based on coccoliths (CÊPEK in Fig. 1. Piece of Mexican amber containing fi ve complete or partial specimens of Termitaradus protera POINAR & DOYEN (white arrows show four specimens), together with seven complete or partial worker termites (black arrows show two specimens). – Scale: 4.0 mm. POINAR & HEISS, HEMIPTERA IN MEXICAN AND DOMINICAN AMBER 53 SCHLEE 1990). Most of the amber is secondarily depos- (Fig. 2). Specimen No. 2, a female, has only the dorsal side ited in turbiditic sandstones of the Upper Eocene to Lower visible. It is 6.5 mm in length, 3.5 mm in width and also Miocene Mamey Group (DRAPER et al. 1994), so the amber has 14 somatic lobes. Specimen No. 3, a female, is 6.0 mm could be older than the Miocene dates. The Dominican in length and 4.4 mm in width. It is only partially visi- amber specimens are deposited in the POINAR collection ble dorsally and the number of somatic lobes could not be maintained at Oregon State University. determined. Specimen No. 4 had only part of the ventrum visible and specimen No. 5 had only the tip of the abdo- men preserved. Neither sex nor stage could be determined 3. Systematic palaeontology in the last two specimens. It was only possible to count the number of lobules Order Hemiptera on all somatic lobes on one side of specimen No. 1. The Family Termitaphididae head has two lobes, lobe one has 16 lobules and lobe two Genus Termitaradus MEYERS 6 lobules. The prothorax has a single lobe, which has 12 lobules. The mesothorax has two lobes, the fi rst has 9 lob- Termitaradus protera POINAR & DOYEN, 1992 ules and the second 7 lobules. The metathorax has a sin- Figs. 1–2 gle lobe with 8 lobules. The abdomen has 8 lobes with the following number of lobules: the fi rst lobe has 12, second Five specimens of T. protera occur in a single piece lobe 12, third lobe 12, fourth lobe 12, fi fth lobe 12, sixth of Mexican amber. Specimen No. 1, a female, is only vis- lobe 12, seventh lobe 10 and the eighth (last) lobe 4. The ible ventrally. It is 6.4 mm in length, 4.6 mm in width, has number of lobules on the lobes of this specimen is almost 14 somatic lobes and exposed antennae, labium and legs identical to that recorded earlier for T. protera (POINAR & DOYEN 1992). A certain degree of variability occurs with the number of lobules on termitaradids, although number of lobules on the terminal (8th) abdominal lobe