Short communicationA primitive triatomine bug, Paleotriatoma metaxytaxa gen. et sp. nov. (Hemiptera: Reduviidae: Triatominae), in mid-Cretaceous amber from northern Myanmar
Introduction
The family Reduviidae (Hemiptera: Heteroptera), commonly called assassin bugs, is one of the most diverse groups of true bugs. While the members of most subfamilies attack only invertebrates, those of the subfamily Triatominae, often called kissing bugs, are notorious vertebrate feeders and many of the 147 known species vector pathogenic trypanosomes (Lent and Wygodzinsky, 1979, Galvão et al., 2003, Patterson and Guhl, 2010, Otálora-Luna et al., 2015, Galvão and Justi, 2018).
The hosts of triatomines are quite diverse and include mammals, birds, amphibians, reptiles and insects (Rabinovich et al., 2011). Since the Triatominae probably evolved from the Reduviinae, feeding on insects can be considered a primitive feature and the early triatomines probably fed on both insect and vertebrates (Weirauch and Munro, 2009). Likewise, the early vertebrate feeders would have been transitory forms that possessed morphological features of both the Triatominae and Reduviinae.
The present study describes an adult kissing bug in Burmese amber that possesses features of extant members of the Triatominae as well as some features of the Reduviinae. The specimen is considered an intermediate fossil representing an early progenitor of the Triatominae. The only previous triatomine fossils are species of Triatoma Laporte and Panstrongylus Burg in mid-Tertiary amber from the Dominican Republic (Poinar, 2005, Poinar, 2013).
Most of the five tribes of the Triatominae (Alberproseniini, Bolboderini, Cavernicolini, Rhodniini and Triatomini) are mainly American in origin with only a few endemic species occurring in Africa, India and Australia. Diversification of the subfamily in South America was considered to have been due to changes resulting from the Andean uplift and variations in sea levels in North America (Justi et al., 2016). Based on evidence that Burmese amber fossils originated in Gondwana and rafted to SE Asia in the Cretaceous (Poinar, 2018), it is proposed that triatomines evolved in Gondwana. This scenario is congruent with the present distribution of the subfamily with representatives in South America, Africa, India and Australia.
Section snippets
Materials and methods
The specimen originated from the Noije Bum 2001 Summit Site mine excavated in the Hukawng Valley and located southwest of Maingkhwan in Kachin State (26°20′N, 96°36′E) in Myanmar (Cruickshank and Ko, 2003, figs. 1–3). Based on paleontological evidence, this site was dated to the late Albian of the Early Cretaceous (Cruickshank and Ko, 2003), placing the age at 97 to 110 Ma. A more recent study using U-Pb zircon dating determined the age to be 98.79 ± 0.62 Ma or at the Albian/Cenomanian boundary
Systematic paleontology
Characters presented by Schuh and Slater, 1995, Weirauch et al., 2014 and Lent and Wygodzinsky (1979) were used to align the fossil with the Triatominae. These characters include: head lacking a transverse constriction behind the eyes, presence of ocelli on oblique cephalic elevations, antennae inserted on tubercles on lateral side of head, antennal segments 3 and 4 inserted on the apex of 2 and 3, respectively, antennal segment 2 with trichobothria, non-raptorial forelegs, membrane with closed
Discussion
Paleotriatoma metaxytaxa gen. et sp. nov. is regarded as an intermediate fossil bridging the gap between the invertebrate predatory Reduviinae and the vertebrate feeding Triatominae. Intermediate fossils normally contain both plesiomorphic and derived features. An example is the Burmese amber bee, Melittosphex burmensis Poinar and Danforth, 2006, which has some bee characters, but also two ancestral wasp features (Danforth and Poinar, 2011). The presence of ocelli, 3-segmented tarsi, 2 closed
Conclusions
Evidence is provided why Paleotriatoma metaxytaxa gen. et sp. nov. is considered to be an intermediate fossil between the Triatominae and the Reduviinae. While containing some features of the Reduviinae, many characters of Paleotriatoma metaxytaxa gen. et sp. nov. occur in members of the subfamily Triatominae (Lent and Wygodzinsky, 1979, Schuh and Slater, 1995, Weirauch et al., 2014). Based on the fossil and present distribution of the Triatominae, it is proposed that this subfamily evolved in
Acknowledgments
The author thanks Alex Brown for donating the specimen to the Poinar amber collection, Eric Vanderduys for taking the photograph in Fig. 7B, Geoff Montheith for supplying the photograph and literature on Triatoma leopoldi, and Roberta Poinar for reviewing earlier drafts of the manuscript. Thanks are also extended to the reviewers whose comments greatly improved the paper.
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