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The Middle Triassic Lagerstätte of Monte San Giorgio Reveals the Oldest Lace Bugs (Hemiptera: Tingidae): Archetingis Ladinica Gen

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The Middle Triassic Lagerstätte of Monte San Giorgio Reveals the Oldest Lace Bugs (Hemiptera: Tingidae): Archetingis Ladinica Gen Rivista Italiana di Paleontologia e Stratigrafia (Research in Paleontology and Stratigraphy) vol. 124(1): 35-44. March 2018 THE MIDDLE TRIASSIC LAGERSTÄttE OF MONTE SAN GIORGIO REVEALS THE OLDEST LACE BUGS (HEMIPTERA: TINGIDAE): ARCHETINGIS LADINICA GEN. N. SP. N. MATTEO MONTAGNA1, LAURA STRADA2, PARIDE DIOLI3 & ANDREA TINTORI4 1Dipartimento di Scienze Agrarie e Ambientali - Produzione, Territorio, Agroenergia, Università degli Studi di Milano, Via Celoria 2, I-20133 Milano, Italy. E-mail: [email protected] 2Dipartimento di Scienze della Terra “Ardito Desio” - Università degli Studi di Milano, Via Mangiagalli 34, I-20133 Milano, Italy. E-mail: [email protected] 3Museo Civico di Storia Naturale di Milano, Corso Venezia 55, I-20121 Milano, Italy. E-mail: [email protected] 4Dipartimento di Scienze della Terra “Ardito Desio” - Università degli Studi di Milano, Via Mangiagalli 34, I-20133 Milano, Italy. E-mail: andrea. [email protected] To cite this article: Montagna M., Strada L., Dioli P. & Tintori A. (2018) - The Middle Triassic lagerstätten of Monte San Giorgio reveals the oldest lace bugs (Hemiptera: Tingidae): Archetingis ladinica gen. n. sp. n. Riv. It. Paleontol. Strat., 124(1): 35-44. Keywords: Fossil insects; new taxa; Hemiptera; Tingidae; true bugs; Middle Triassic. Abstract. A new genus and species of fossil lace bugs (Hemiptera; Tingidae), Archetingis ladinica gen. n. et sp. n. is described from the Lower Kalkschieferzone (Meride Limestone, Upper Ladinian) of the Swiss side of Monte San Giorgio. The new taxon clearly resembles modern Tingidae in its hemelytra and pronotum, with two rows of areole, in the presence of bucculae closed at the anterior end, not extending beyond the apex of the head, and in the presence of a labial groove on the meso-/metathoracic sternum. Distinctive features are the thick femurs of the first and second pair of legs, the exceptional size (total length of ~12 mm) if compared with both fossil and extant species. Archetingis ladinica gen. n. et sp. n. represents by far the oldest known species of this family and brings back the origin of Tingidae of approximately 140 My, well into the Middle Triassic. The discovery of A. ladinica, beside its evolutionary consequences on the origin of extant Tingidae, provides evidences for the presence of terrestrial ecosystems nearby to the depositional environment. According to the living and trophic behavior of extant Tingi- dae, those emerged lands had to be covered by vegetation. INTRODUCTION of these rarely preserved groups of organisms. In spite of the detailed scientific excavations carried Monte San Giorgio (MSG, Italy-Switzerland) out for more than 100 years, the fossil invertebrates is a Triassic Fossil-Lagerstätte inscribed in the have been discovered only recently (Lombardo et UNESCO World Heritage List for its marine ver- al. 1999). tebrate faunas, (Felber et al. 2004; Felber 2005). Pa- During the fieldwork carried out between leontological researches on MSG date back to the 1997 and 2003 in the Lower Kalkschieferzone (Late middle of XIX Century, when the first specimens Ladinian, 239.51 ± 0.15 Ma; Stockar et al. 2012) on were discovered on the Italian side of the moun- the Swiss side of Monte San Giorgio, a remarkably tain. The unique aspect of MSG is that it concen- diverse assemblage of 19 insect specimens were col- trates several different fossil vertebrate horizons lected. The specimens were attributed to six orders in a few square km (Felber et al. 2004) covering a and to groups characterized by different ecology and time interval from Late Anisian to Late Ladinian. life habits (Strada et al. 2014). So far, seven speci- The scientific interest on MSG has been almost al- mens from this assemblage, ascribed to four taxa, ways focused on marine vertebrates or on inverte- have been formally described: the mayfly Tintorina brates dealing with biostratigraphy. More recently, meridensis Krzeminski & Lombardo, 2001, an elytron the invertebrates collected from three levels of of the beetle Notocupes sp. (Krzeminski & Lombar- MSG lagerstätte, among which insects, proved to do 2001), the beetle Praedodromeus sangiorgensis Strada be of relevant meaning for both paleoenvironmen- et al., 2014 (see also Krzeminski & Lombardo 2001) tal understanding and evolutionary reconstruction and a representative of extant bristletail Gigamachil- is triassicus Montagna et al., 2017. Noteworthy, the Received: June 26, 2017; accepted: October 26, 2017 entomofauna from the Middle Triassic of Monte 36 Montagna M., Strada L., Dioli P. & Tintori A. Fig. 1 - Geographic position of the Monte San Giorgio (Italy-Switzerland) UNESCO World Heritage, with the strata of the Lower Kalkschie- ferzone (Site D Val Mara) excavation from which the specimens were collected (left; indicated by white arrows: Archetingis ladinica collected from stratum 18 of site D, Val Mara; remnant of Voltzia collected on stratum 21 of site D, Val Mara; the vertical black bar corresponds to 1 meter) and stratigraphic section of the Middle Triassic sediments in the Monte San Giorgio with distribution of the insect fossils (right). Stratigraphic log modified from Bechly & Stockar (2011). San Giorgio includes terrestrial groups, with phy- herein described and named. tophagous and predatory habits, as well as aquatic Within true bugs, Cimicomorpha, with ap- groups, collected as larvae (e.g., Plecoptera) and as proximately 20.000 described species, represent adults (e.g., Ephemeroptera). The diversity of the the most diverse and successful lineage of Het- entomofauna from the Kalkschieferzone, together eroptera. Representatives of this group possess with the high biodiversity of the fish assemblage heterogeneous ecology and have successfully (Lombardo et al. 2012), supports the presence of colonized a broad variety of habitats, with the a complex paleoenvironment in the area of Monte evolution of different habits spanning from pre- San Giorgio, which is known to have been a shal- dation to phytophagy, the former retained as the low lagoon, with limited connections to the open groundplan feature of Heteroptera (Cobben 1978; and deeper sea, adjacent to a carbonate platform, Schuh et al. 2009). On the basis of the fossil re- with permanent fresh water pools or small streams cord, the first extant cimicomorphan families have (Strada et al. 2014). Remarkably, so far no amphib- made their appearance in the Jurassic, with the first ian remains have been collected, although the pa- occurrence represented by Nabidae (Cimicoidea) leoenvironment looks favourable for their life and and Miridae (Miroidea) in the Jurassic (Carpen- they are quite common in the coeval rocks of the ter 1992; Hou et al. 2012; Nicholson et al. 2015). German Basin (Schoch 2006). Within Tingoidea, three extant families have been Among the insect fossil assemblage of recognized by Lis (1999), namely the Vianaidi- Monte San Giorgio one specimen attributed to dae, Tingidae, and Cantacaderidae. Other authors true bugs was discovered (Strada et al. 2014) and is (Froeschner 1996; Froeschner 2001; Golub 2001) The oldest lace bugs (Hemiptera: Tingidae) 37 recognised only the two families of Vianaididae were processed using Adobe Photoshop CS3. The drawing of the ® and Tingidae, the latter including Cantacaderinae taxon has been produced using graphics tablet (Intuos pro, Wa- com) and Adobe Illustrator CC. as a subfamily together with Tinginae (Schuh et al. 2006). Members of Vianaididae and Tingidae have been recovered from the Cretaceous Peri- TAPHONOmiC REMARKS od (Popov 1989; Golub & Popov 2000; Golub & Popov 2003a; Perrichot et al. 2006; Golub & The specimen here described, preserved in Popov 2008). In addition to Vianaididae and Tin- part and counterpart, has maintained the general gidae, three new fossil tingoid families have been layout of the body, including the three tagma, as recently described, namely Ignotingidae from the well as most of the appendages. However, two out Laiyang Formation at the Jurassic-Cretaceous of the six legs and the two antennae, with the ex- boundary (Zhang et al. 2005), Ebboidae from Al- ception of the basal antennomeres, are displaced. bian amber of France (Perrichot et al. 2006) and This preservation suggests that the organism went Hispanocaderidae from Lower Cretaceous amber through a first stage of decay in subaerial and/or of Spain (Golub et al. 2012). superficial aquatic environment, during which part Within Tingidae, encompassing approxi- of appendages were disarticulated and lost. There- mately 2100 extant species worldwide distributed fore, we hypothesize that short after the death and (Froeschner 1996), more than 40 fossil species a preliminary decay on terrestrial environment, the are known from Lower Cretaceous to the Upper insect was transported by not high-energy events, Miocene (Golub 2001; Wappler 2003; Golub & such as a water run-off after monsoons (as pre- Popov 2003a; Golub & Popov 2003b; Nel et al. viously suggested, see Tintori 1990a,b; Tintori & 2004; Perrichot et al. 2006; Wappler 2006; Golub Brambilla 1991) or by wind, to the depositional & Popov 2008; Heiss & Guilbert 2013; Wappler et basin where, after a fast sinking to the bottom, al. 2015). Thus, currently the oldest lace bugs fos- burial and fossilization took place. Since most of sils are Sinaldocader drakei Popov, 1989 and Sinaldo- the specimen’s body layout is preserved, we can cader ponomarenkoi Golub & Popov, 2008 from the hypothesize a time-limited decay, a short transpor- Lower Cretaceous of Eastern Siberia, and
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