Cariridris bipetiolata

†Cariridris bipetiolata Temporal range: Aptian, Cretaceous K Pa Eo Ol Mi Pl Crato, Brazil
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Hymenoptera
Family:	Sphecidae
Genus:	Cariridris
Species:	†C. bipetiolata
Binomial name
†Cariridris bipetiolata Brandão & Martins-Neto, 1990

This species has been transferred to the family Sphecidae, possibly in the (sub)family Ampulicidae.

A real surprise was the record of a Formicidae from the Lower Cretaceous of the Crato Formation (Brandao et al., 1989). However, the attribution of the specimen was at first not very confident at all, and is still disputed to this day (J. Rust, personal communication). The preservation of the holotype is relatively poor, with indistinct edges and incomplete cephalic area. Furthermore, it is not possible to see whether the specimen had wings or not. Subsequently, the animal has been mistaken for a representative of the Ampulicidae (‘Sphecidae’), which often have an extremely similar general habitus (Figure 11.72e). However, the structure of the petiolus indicates that the fossil indeed belongs to the Formicidae of the subfamily Myrmeciinae (bulldog ants), whose distribution is now limited to Australia. As it was not possible to attribute the new fossil species to the single Recent genus Myrmecia, it was named Cariridris bipetiolata by Brandao et al. (1989) (Osten, 2007).

Dlussky & Rasnitsyn (2003) treated this species as belonging to the family Sphecidae rather than Formicidae. They state: C. bipetiolata as described and figured by Brandão et al. (1989) looks very similar to an unnamed Santana fossil kept at the American Museum Natural History (New York, New York, USA) and described as belonging to either Sphecidae (Ampulicinae) or Dryinidae (Darling, Sharkey, 1990). One of us (APR) jointly with W.J. Pulawski of the California Academy of Sciences (San Francisco, California, USA) have studied that fossil (AMNH 44108) and found it belonging to Sphecidae (probably to subfamily Ampulicinae) rather than to Formicidae. That is why we maintain here our earlier conclusion that Cariridris is a genus of Sphecidae (Rasnitsyn, 1990; Dlussky, 1998).

Two further specimens of Cariridris may have been discovered while examining fossil material by Osten (2007). There is no doubt that these two fossils also belong to the same genus and species Cariridris bipetiolata (Figure 11.77). The specimen SMNS 66565 (Plate 15m) is much better preserved than the holotype, and supports the attribution to Formicidae–Myrmeciinae (petiolus and head structures). The absence of wings in this specimen is of course no valid evidence against such an attribution.

Dlussky (2012) followed the treatment of Dlussky and Rasnitsyn (2003) and considered this species to be incertae sedis in Ampulicidae. However, it appears he was unaware of the specimens examined by Osten (2007) (as he does not cite this paper).

Jouault & Nel (2021) note that Osten (2007) lacked justification and that no illustrations were provided to support the placement of this taxon within the ants. Because of this, they did not consider this taxon to belong to the family Formicidae, but did not speculate on its proper placement.

Distribution

This taxon was described from Crato, Brazil (Aptian, Cretaceous).

Nomenclature

The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.

• †bipetiolata. †Cariridris bipetiolata Brandão & Martins-Neto, in Brandão, et al. 1990: 201, figs. 1-3 (q.) BRAZIL (Ceará, Cretaceous).
  • Status as species: Brandão, 1990a: 313; Brandão, 1991: 337; Bolton, 1995b: 134.
  • Excluded from Formicidae, incertae sedis in ACULEATA: Grimaldi, Agosti & Carpenter, 1997: 7; Ward & Brady, 2003: 362.
  • Excluded from Formicidae, incertae sedis in “sphecid wasp subfamily” Ampulicinae: Rasnitsyn, 2002: 249; Dlussky & Rasnitsyn, 2002: 418; Dlussky, 2012: 288 (in text).
  • In Formicidae, Myrmeciinae: Osten, 2007: 353.
  • Excluded from Formicidae, transferred to Sphecidae: Jouault & Nel, 2021: 1, 5 (exact placement not specified).

References

• Barden, P. 2017. Fossil ants (Hymenoptera: Formicidae): ancient diversity and the rise of modern lineages. Myrmecological News 24: 1-30.
• Bolton, B. 1995b. A new general catalogue of the ants of the world. Cambridge, Mass.: Harvard University Press, 504 pp. (page 134, Cariridris incertae sedis in Myrmeciinae)
• Borowiec, M.L., Moreau, C.S., Rabeling, C. 2020. Ants: Phylogeny and Classification. In: C. Starr (ed.), Encyclopedia of Social Insects (doi:10.1007/978-3-319-90306-4_155-1).
• Brandão, C.R.F., Martins-Neto, R.G. & Vulcano, M.A. 1990. The earliest known fossil ant (first southern hemisphere Mesozoic record). Psyche 96: 195-208.
• Dlussky G.M. 1998. [Formicoidea of Cretaceous and age of ants]. Materialy X Symposiuma "Muravji i zaschita lesa". Moscow.P.5-8 [in Russian].
• Dlussky, G.M. 2012. New fossil ants of the subfamily Myrmeciinae from Germany. Paleontological Journal 46 (3): 288-292.
• Dlussky, G.M., Rasnitsyn, A.P. 2003. Ants (Hymenoptera: Formicidae) of Formation Green River and some other Middle Eocene deposits of North America. Russian Entomological Journal 11(4): 411-436.
• Fernandez, F., Guerrero, R.J., Sánchez-Restrepo, A.F. 2021. Sistemática y diversidad de las hormigas neotropicales. Revista Colombiana de Entomología 47, 1–20 (doi:10.25100/socolen.v47i1.11082).
• Grimaldi, D.; Agosti, D.; Carpenter, J. M. 1997. New and rediscovered primitive ants (Hymenoptera: Formicidae) in Cretaceous amber from New Jersey, and their phylogenetic relationships. American Museum Novitates 3208: 1-43 (page 7, Cariridris incertae sedis in Aculeata)
• Jouault, C., Nel, A. 2021. The oldest Cenozoic ant fossil: †Tyrannomecia gen. nov. (Formicidae: Myrmeciinae) from the Palaeocene Menat Formation (France). Historical Biology, 1–8 (doi:10.1080/08912963.2021.2010192).
• Osten, T. 2007. 11.18 Hymenoptera: bees, wasps and ants. Pp. 350-365 in: Martill, D. M., Bechly, G., Loveridge R. F. (eds.) 2007. The Crato Fossil Beds of Brazil: Window into an Ancient World. Cambridge: Cambridge University Press, 625 pp. (doi:10.1017/CBO9780511535512).
• Rasnitsyn A.P.1990. Table 1. Known Cretaceous Hymenoptera, in Darling D.Ch., Sharkey M.J., Order Hymenoptera, in D. A. Grimaldi (ed.): Insects from the Santana Formation, Lower Cretaceous, of Brazil. Bull. Amer. Mus. Nat. Hist. 195: 124-129.
• Ward, P., and Brady, S. 2003. Phylogeny and biogeography of the ant subfamily Myrmeciinae (Hymenoptera: Formicidae). Invertebrate Systematics 17(3): 361-386 (page 362, Cariridris incertae sedis in Aculeata)