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Photocryptus concinnus (Brullé) (Photo by BB Longo)
Photocryptus concinnus (Brullé)
(BB Longo photo)

Bernardo F. Santos, Hymenoptera
Peter Buck Postdoctoral Fellow

  • Phone:   202-633-1003
  • Fax:   202-633-4565
  • E-mail Address:
  • USPS Address:
    Smithsonian Institution
    PO Box 37012, MRC 188
    Washington, DC 20013-7012
  • Public Carrier Address:
    Smithsonian Institution
    National Museum of Natural History
    10th & Constitution NW
    Washington, DC 20560-0188
  • Education:

    Ph.D. in Comparative Biology.
    Richard Gilder Graduate School
    (American Museum of Natural History)

    B.S. in Biology, M.Sc. in Animal Biology. 
    Universidade Federal do Espírito Santo, Brazil.
Bernardo F. Santos

Research Interests:

My research interests are centered on the evolution of one of the most diverse and ecologically important groups of organisms on Earth: the ichneumonid parasitic wasps (Hymenoptera, Ichneumonidae). I am particularly interested in the processes that generated the impressive diversity of life history strategies, morphological adaptations and ecological traits seen in these wasps. During my Ph.D., I have worked to elucidate these questions by combining a robust phylogenetic framework with statistical tools such as phylogenetic comparative methods and geometric morphometrics. In the process of investigating these issues, I also invest in unraveling the largely unexplored wasp biodiversity, conducting projects of revisionary systematics and describing new species.

Systematics of Cryptine wasps:

This is my primary and most basic line of research. Since the beginning of my career, I have been dedicated to understand the evolutionary history of Cryptini and describe and classify the biodiversity of this hyperdiverse group of wasps. Since 2008, I have described 92 new species and one new genus, and many more new species are identified and waiting to be described! During my Ph.D. I have also produced a major phylogenetic study, the first to investigate in detail the evolutionary history of Cryptini. This work serves as a basis to the broader evolutionary and ecological aspects of my research.

Morphological diversification

At present, one of my main research interests is the study of phenotypic adaptations, particularly convergent evolution. I have been focusing on the evolutionary pathway of adaptations related to finding and assessing deeply concealed hosts by cryptine wasps. I am using comparative methods and geometric morphometrics in a phylomorphospace approach to investigate the idea that these adaptations are part of a complex morpho-functional system, and how that affects the evolution of body shape in the associated lineages.

Other ongoing projects and interests include:

Color patterns. Cryptines display an impressive variety of color patterns, which seem to show a pattern of convergence across lineages. I am investigating the phylogenetic and geographic signals shown in color patterns in a group of cryptine wasps and what these patterns can tell us about evolutionary processes.

Species delimitation. In partnership with my outstanding high school mentees, I am using molecular markers and geometric morphometrics to investigate species limits in a genus of Andean cryptine wasps, Cryptopteryx. Preliminary results indicate that several cryptic species may be present in this widely distributed group.

Surveys and inventories. Only by continuously documenting species occurrence we can draw a truly informative picture of Earth’s biodiversity and understand which environmental factors help to shape communities and ecosystems. In partnership with colleagues, I am involved in several surveys and community ecology studies throughout the Neotropics.

Publications   google scholar

Santos, B.F. Accepted. Phylogeny and reclassification of Cryptini (Hymenoptera, Ichneumonidae, Cryptinae), with implications for ichneumonid higher-level classification. Systematic Entomology.

Santos, B.F., Aguiar, A.P., Tedesco, A.M. & Fontenelle, J.C.R. 2017. Long-term seasonal dominance of the wasp Trihapsis polita Townes (Hymenoptera, Ichneumonidae) in the Brazilian Atlantic Forest. Biodiversity Data Journal, 5, e11337.

Ceríaco, L.M.P., Gutierrez, E., Dubois, A. et al., Santos, B.F. et al. 2016. Photography-based taxonomy is inadequate, unnecessary, and potentially harmful for biological sciences. Zootaxa (Auckland. Print), 4196: 435-445.PDF

Santos, B.F. 2016. Generic redefinition and a new species of Nesolinoceras (Hymenoptera, Ichneumonidae, Cryptinae). American Museum Novitates, 3858: 1–16.

Villanueva-Bonilla, G.A., Onody, H.C., Santos, B.F. & Vasconcelos-Neto, J. 2016. First record of egg sac predation on a wall crab spider, Selenopidae (Araneae), by the wasp Camera lunavenatrix n. (Ichneumonidae, Cryptinae). Journal of Hymenoptera Research, 49: 65–79.

Aguiar, A.P. & Santos, B.F. Revision of Melanocryptus Cameron (Ichneumonidae, Cryptinae), with description of seven new species. American Museum Novitates, 3836: 1–56.

Santos, B.F., Payne, A., Pickett, K.M. & Carpenter, J.M. 2015. Phylogeny and historical biogeography of the paper wasp genus Polistes (Hymenoptera: Vespidae): implications for the overwintering hypothesis of social evolution. Cladistics, 31: 535–549.

Santos, B.F.
& Aguiar, A.P. 2015. Review of Loxopus Townes (Hymenoptera, Ichneumonidae, Cryptinae), with descriptions of six new species. Journal of Natural History, 49(31-32): 1905–1935.

Santos, B.F. & Aguiar, A.P. 2013. Phylogeny and revision of Messatoporus Cushman (Hymenoptera, Ichneumonidae, Cryptinae), with description of sixty five new species. Zootaxa, 3634: 1–284.PDF

Aguiar, A.P. & Santos, B.F. 2012. Additions to the revision of Digonocryptus Viereck (Hymenoptera, Ichneumonidae, Cryptinae), with nine new taxa, new males and distribution maps for all known species. Zootaxa, 3282: 1–41.PDF

Santos, B.F.
, Aguiar, A.P. & Tedesco, AM. 2012. Trigonalidae (Hymenoptera) from cacao agroforestry systems in northeastern Brazil, with two new species of Trigonalys. Journal of Hymenoptera Research, 25: 19–33.

Santos, B.F.
& Aguiar, A.P. 2012. Phylogeny and description of Eknomia, a morphologically unusual new genus of Neotropical Cryptinae (Hymenoptera, Ichneumonidae, Cryptinae). Zootaxa, 3237: 35–52.PDF

Aguiar, A.P. & Santos, B.F. 2010. Discovery of potent, unsuspected sampling disparities for Malaise and Möricke traps, as shown for Neotropical Cryptini (Hymenoptera, Ichneumonidae). Journal of Insect Conservation, 14: 199–2010.

Scherrer, M.V., Santos, B.F. & Aguiar, A.P. 2010. Generic redefinition, key and two new species of Acorystus Townes (Hymenoptera, Ichneumonidae, Cryptinae). Zootaxa, 2721: 28–38.PDF

Santos, B.F.
, Aguiar, A.P. & Tedesco, A.M. 2009. Phylogenetic revision and the origin of Polyphrix Townes (Hymenoptera, Ichneumonidae, Cryptinae), with description of a new species. Zootaxa, 2214: 29-44.PDF

Aguiar, A.P. & Santos, B.F. 2009. Biological synopsis of Photocryptus Viereck (Hymenoptera, Ichneumonidae, Cryptinae), with eight new host records and fourteen new geographic records. Zootaxa, 2148: 49–45.PDF

Santos, B.F.
& Aguiar, A.P. 2008. Phylogeny and reclassification of Distictus Townes (Hymenoptera, Ichneumonidae, Cryptinae), with description of a new species. Zootaxa, 1934: 30–39.PDF

Book chapter

Aguiar, A.P., Santos, B.F., Couri, M.S., Rafael, J.A., Costa, C. Ide, S., Duarte, M., Grazia, J., Schwertner, C.F., Freitas, A.V.L. & Azevedo, C.O. 2009. Insecta. In: Rocha, R.M. & Boeger, W.A. Estado da Arte e Perspectivas para a Zoologia no Brasil [State of Art and Perspectives to the Zoology in Brazil]. Editora UFPR, Curitiba, pp.131–155.


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