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Smithsonian National Museum of Natural History

Department of Paleobiology

  • Jorge A. Santiago-Blay
  • Resident Research Associate
  • Phone:   202-633-1383
  • Fax:   202-786-2832
  • E-mail Address:   blayj
  • Mailing Address:
    Smithsonian Institution
    PO Box 37012, MRC 121
    Washington, DC 20013-7012
  • Shipping Address:
    Smithsonian Institution
    National Museum of Natural History
    10th & Constitution NW
    Washington, DC 20560-0121

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Ph.D. University of California, Berkeley. Entomological Sciences. 1990
M.A. University of California, Berkeley. Botany. December 1989.
M.S. University of Puerto Rico, Río Piedras. Biology. 1985.
B.S. University of Puerto Rico, Río Piedras. Biology. 1979. Minors: Mathematics and Education. Magna cum Laude.

Research Interests

The main focus of my research seeks to provide answers to this question: What processes cause and what patterns correlate with speciation and biodiversity? Because my training, experience, and interests are multidisciplinary, I seek answers from numerous areas of biology, including paleobiology, ecology, behavior, genetics, biochemistry, and others using different biological systems. Therefore, I can comfortably accommodate those who wish to do research with me on a wide variety of topics of mutual interest particularly, but not exclusively, arthropods and plants.

I have had the pleasure of collaborating with numerous people engaged in research projects. These projects include subjects as diverse as: plant developmental genetics, insect cytogenetics, aquatic ecology, reconstruction of ancient communities preserved in the fossil record, forensic ecology, environmental ethics, biological illustration, biology education, and many others. In collaboration with each person, some of them students, research initiatives are written and reviewed. Hence, competitive grant proposals may be submitted and progress reports presented at professional meetings. Some have received funding to pursue their research projects. As a result of their increasing intellectual independence and long-term productivity, I have more time to pursue other research interests (as seen here, on page 25.).  Below, I briefly summarize some of my research interests, in no particular order.

Paleobiology. The natural history, biogeography, and ecology of ancient communities. For instance, the study of two North American Jurassic assemblages preserved in lithographic limestone and seemingly dominated by insects suggests that it is ecologically and biogeographically closer to its penecontemporaneous counterparts in Asia and not those described for Meso- and South America. Also, there are great differences in the botanical composition between these two North American Jurassic paleocommunities. We have conducted similar research on ancient communities preserved in amber collected in different parts of the world.

Plant exudate chemistry. Our ultimate goal is to generate widely available databases of C-13 (carbon-13 isotope) solid state nuclear magnetic resonance spectroscopy and H-1 (proton) NMR signatures of plant exudates of the world. With a larger database, we will have greater ability to: 1) identify the botanical origin of modern and fossil exudates (e.g. amber), 2) make specific statements about the systematics and evolution of some exudate-producing plant groups (e.g. Fabaceae), and 3) understand paleoenvironments and paleoclimates entombed in fossil exudates during different intervals in Earth's history.

Aquatic Ecology. I am especially interested in documenting stream biodiversity and ascertaining the water quality therein. In a 13 month-long study done with undergraduates, we showed a discrete gap in the relative composition of biodiversity along Snyder’s Branch Creek (Salem, VA). The nature of this gap suggested the presence of an organic pollution source. Furthermore, we documented a large biodiversity of aquatic macroinvertebrates, including over 50 species of larval chironomids.

Sexual Selection and Evolution. What makes Drosophila melanogaster females choose conspecific males? The answer to that question involves many sensory channels whereby females obtain information about the conspecificity and possibly “quality” of courting males. We have shown that while female D. melanogaster prefer fully winged over wingless males, they cannot discriminate between fully winged males and males with over one complete wing and three-quarters, or more, of another wing.

Molecular identification and phylogenetics of arthropods. Our investigations have confirmed the usefulness of molecular markers for the rapid and reliable identification of agriculturally, silviculturally, and medically important insects, including the agriculturally-beneficial parasitic wasps, Trichogramma, the cone-feeding beetles, Conophthorus, and the blood sucking sand fly Lutzomyia. For years, several colleagues and I have been working on the large Puerto Rican arboreal millipedes Anadenobolus arboreus.

Insect-plant interactions: a case of possible coevolution between species of North American Monoxia beetles (Chrysomelidae) and their host plants. Monoxia is especially interesting because most of its species groups feed on selected host plant genera placed in very divergent plant families, the Asteraceae, the daisies, and the Chenopodiaceae, the plants in the spinach family. By doing field and laboratory work, I have found congruent patterns of host plant association, cytogenetics, and morphology.

For additional information, please feel free to visit my personal website

Current Editorial Service

  • Editor-in-Chief, Life: The Excitement of Biology
  • Scientific Co-Editor, Research on Chrysomelidae
  • Associate Editor, Journal of Agriculture of the University of Puerto Rico

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Research papers in refereed journals

1. Santiago-Blay, J. A. and M. J. Vé1ez. 1980. Nota sobre una posible nueva especie y un nuevo récord de diplópodos de Puerto Rico. Science-Ciencia. Boletín Científico del Sur (Ponce, Puerto Rico) 7:54-55.

2. Santiago-Blay, J. A. 1983. Annotated list of the scorpion taxa reported for Puerto Rico and adjacent islands. Science-Ciencia. Boletín Científico del Sur (Ponce, Puerto Rico) 10:92-93.

3. Santiago-Blay, J. A. 1983. Plumeria rubra: A new host plant record for the Fall Armyworm (Lepidoptera: Noctuidae) in Puerto Rico. Florida Entomologist 66:359.

4. Francke, O. F. and J. A. Santiago-Blay. 1984. Redescription of Tityus crassimanus (Thorell, 1877) and its junior synonym Tityus antillanus (Thorell, 1877) (Scorpiones: Buthidae). Journal of Arachnology 12:283-290.

5. Santiago-Blay, J. A. 1985. Microtityus dominicanensis: A new buthid scorpion from Hispaniola, West Indies. Entomological News 96:1-6.

6. Santiago-Blay, J. A. 1985. Redescription of Tityus dasyurus dasyurus Pocock, 1897 (Scorpiones: Buthidae). Revue Arachnologique 6:49-56.

7. Santiago-Blay, J. A. 1985. Aclaraciones en torno a los significantes zoológicos de la voz "alacrán" en Puerto Rico. Science-Ciencia. Boletín Científico del Sur (Ponce, Puerto Rico) 12:43-45.

8. Santiago-Blay, J. A. and M. J. Vélez, Jr. 1985. Chondromorpha xanthotricha: A new record of polydesmid millipede (Diplopoda: Paradoxosomatidae) and the geographic distribution of the three known paradoxosomatids of Puerto Rico. Caribbean Journal of  Science 21:137-141.

9. Santiago-Blay, J. A. 1985. Notes on Pseudosphinx tetrio (L.) (Lepidoptera: Sphingidae) in Puerto Rico. Journal of the Lepidopterist’s Society 39:208-214.

10. Santiago-Blay, J. A. 1986. Morphological malformations among scorpions of Puerto Rico and the adjacent islands. Pan-Pacific Entomologist 62:77-82.

11. Santiago-Blay, J. A. and S. Medina Gaud. 1986. Notes on the biology of Aspisoma ignitum: A new firefly for Puerto Rico (Coleoptera: Lampyridae). Florida Entomologist 69:440-444.

12. Casas-Benabe, R. and J. A. Santiago-Blay. 1987 ["1986"]. First case report of scorpion sting in Puerto Rico. Caribbean Journal of Science 22:213-214.

13. Santiago-Blay, J. A. 1987. The scorpions of Dominica (West Indies). Journal of Entomological Science 22:311-316. (Erratum 1988. 23:96.)

14. Santiago-Blay, J. A. 1987. Medical importance of scorpion stings in Puerto Rico: Preliminary Survey. Caribbean Journal of Science 23:314-317.

15. Santiago-Blay, J. A. and J. Maldonado-Capriles. 1988. Observations on the true bugs Emesa tenerrima, a possible spider mimic, and Ghinallelia borincana (Hemiptera: Reduviidae: Emesinae) from Puerto Rico. Proceedings of the  Entomological  Society of Washington 90:369-372.

16. Santiago-Blay, J. A. and G. O Poinar, Jr. 1988. A fossil scorpion, Tityus geratus new species (Scorpiones: Buthidae) from Dominican amber. Historical Biology 1:345-354.

17. Santiago-Blay, J. A. and H. V. Daly. 1989 ["1988"]. The Apoidea collection of the California Insect Survey (C.I.S.) at the Essig Museum of Entomology, University of California, Berkeley. The Pan-Pacific Entomologist 64:281-284.

18. Santiago-Blay, J. A. and D. L. Pardoe. 1989. The spiny stub: an alternative technique for mounting specimens for SEM. Bulletin of the Electron Microscopy Society of America 19:112.

19. Poinar, G. O., Jr. and J. A. Santiago-Blay. 1989. A fossil solpugid, Happlodontus proterus, new genus, new species (Arachnida: Solpugida) from Dominican amber. Journal of the New York Entomological Society 97:125-132.

20. Santiago-Blay, J. A. and N. Virkki. 1989. On the XO sex chromosome system of Aulacoscelis melanocera Stål (Aulacoscelinae: Chrysomelidae: Coleoptera). Hereditas 111:99-102. (Erratum:  Duponchel and Chevrolat, not Stål described, this species.)

21. Santiago-Blay, J. A. 1989. Chalcidoids (Hymenoptera) reared from Artemisia tridentata (Asteraceae) galls in British Columbia, Canada. Journal of the Entomological Society of British Columbia  86:80-81. (Errata  1991. 87:90.)

22. Santiago-Blay, J. A. 1990 ["1989"]. Seasonal occurrence and host plant feeding preferences of adult Monoxia n. sp. 1 (Coleoptera: Chrysomelidae: Galerucinae). Entomography 6:397-401.

23. Santiago-Blay, J. A.  and J. Maldonado Capriles. 1990 ["1989"]. A new structure on the hind legs of male Monalocoris carioca Carvalho and Gomez (Heteroptera: Miridae). Journal of the New York Entomological Society 97: 479-482.

24. Santiago-Blay, J. A., W. Schawaller and G. O. Poinar, Jr. 1990. A new specimen of Microtityus ambarensis (Schawaller, 1982) (Scorpiones: Buthidae), fossil scorpion from Hispaniola evidence of the taxonomic status and possible biogeographic implications. Journal of Arachnology 18:115-117.

25. Maldonado Capriles, J. and J. A. Santiago-Blay. 1991. Classification of Homalocoris (Heteroptera: Reduviidae: Hammacerinae), with the description of a new species. Proceedings of the Entomological Society of Washington 93:703-708.

26. Virkki, N., J. A. Santiago-Blay, and E. G. Riley. 1992. Chromosomes of Puerto Rican Hispinae and Cassidinae (Chrysomelidae). Coleopterist’s Bulletin 46:29-42.

27. Santiago-Blay, J. A. and G. O. Poinar, Jr. 1992. Millipedes from Dominican amber, with the description of two new species of Siphonophora (Diplopoda: Siphonophorida: Siphonophoridae). Annals of the Entomological Society of America 85:363-369.

28. Virkki, N., J. A. Santiago-Blay, and S. M. Clark. 1992 ["1991"]. Chromosomes of some Puerto Rican Disonychina and Oedionychina (Coleoptera: Chrysomelidae: Alticinae: Oedionychini): Evolutionary Implications. Psyche 98:373-390. (Errata 99:140.)

29. Maldonado Capriles, J. and J. A. Santiago-Blay. 1992. A new species of the Neotropical genus Sphaeridops Amyot & Serville, 1843 (Sphaeridopinae: Reduviidae). Proceedings of the Entomological  Society of Washington 94:508-511.

30. Santiago-Blay, J. A. and G. O. Poinar, Jr. 1993. Classification of Diphleps (Heteroptera: Miridae: Isometopinae), with the description of D. yenli, a new species from Dominican amber (lower Oligocene - upper Eocene). Proceedings of the Entomological Society of Washington 95:70-73.

31. Maldonado Capriles, J., J. A. Santiago-Blay, and G. O. Poinar, Jr. 1993. Paleoploiariola venosa, a new fossil Emesinae (Heteroptera: Reduviidae) genus and species from Dominican amber. Journal of Agriculture of the University of Puerto Rico 77:95-100.

32. Santiago-Blay, J. A. and S. Battaion. 1993. Non-nutrient agar: an alternative cheap matrix for electrophoresis. Trends in Genetics 9:158.

33. Santiago-Blay, J. A. 1993. The scorpions of Hispaniola and Puerto Rico. American Arachnology 47:6-7.

34. Maldonado Capriles, J., J. A. Santiago-Blay, and G. O. Poinar, Jr. 1993. Apicrenus fossilis gen. & sp. n. (Heteroptera: Reduviidae: Apiomerinae) from Dominican amber (lower Oligocene – upper Eocene). Entomologica Scandinavica 24:139-142.

35. Fain, A. and J. A. Santiago-Blay. 1993. A new genus and two new species of mites (Acari) phoretic or parasitic on Neotropical Chrysomelidae (Coleoptera). Acarologia 34:337-343.

36. Santiago-Blay, J. A. and G. O. Poinar, Jr. 1993. First scorpion (Buthidae: Centruroides) from Mexican amber (lower Miocene to upper Oligocene). Journal of Arachnology 21:147-151.

37. Virkki, N. and J. A. Santiago-Blay. 1993. Trends of karyotype evolution in Neotropical Oedionychina (Coleoptera: Chrysomelidae: Alticinae). Hereditas 119:263-283.

38. Virkki, N. and J. A. Santiago-Blay. 1995. ["1994"]. Chromosomes of some Neotropical flea beetles (Coleoptera: Chrysomelidae: Alticinae) with inherent univalent, amphiorientating sex chromosomes and meioformula 11+X+y in the male meiosis. Cytobios 80:79-99.

39. Santiago-Blay, J. A. and T. L. Young. 1995. Reliable sexing of adult Conophthorus (Coleoptera: Scolytidae) beetles. Canadian Entomologist 127:605-607.

40. Maldonado Capriles, J., J. A. Santiago-Blay, and G. O. Poinar, Jr. 1996. Paralienates hyalinus n. gen., n. sp. (Enicocephalidae: Heteroptera): A fossil gnat bug from Dominican amber (lower Miocene - upper Eocene). Journal of Agriculture of the University of Puerto Rico 80: 47-54.

41. Virkki, N. and J. A. Santiago-Blay. 1996. Atypical cytology in some Neotropical flea beetles (Coleoptera: Chrysomelidae: Alticinae: Oedionychina) from an intense natural radiation site known, Morro do Ferro (Brazil). Cytobios 85:167-184.

42. Santiago-Blay, J. A. and C. E. Ayala. 1996. A minuten probe for small organisms. Entomological News 107:245-246.

43. Virkki, N. and J. A. Santiago-Blay. 1996. Chromosomes of some Puerto Rican flea beetles (Coleoptera: Chrysomelidae: Alticinae): Multiple cytogenetic evolutionary tendencies in the Neotropics. Journal of Zoological Systematics and Evolutionary Research. Zeitschrift für zoologische Systematik und Evolutionsforschung 34:113-119.

44. Santiago-Blay, J. A. and N. Virkki. 1996. Evolutionary relationships within Monoxia (Coleoptera: Chrysomelidae: Galerucinae): Chromosomal evidence for its intrageneric classification. Caryologia (Firenze, Italy) 49:257-265.

45. Santiago-Blay, J. A. 1996. Morphometrics of two Californian populations of Gumaga nigricula (McLachlan 1871) (Trichoptera: Sericostomatidae). Journal of Agriculture of the University of Puerto Rico 80:157-167.

46. Poinar, Jr. G. O. and J. A. Santiago-Blay. 1997. Paleodoris lattini gen. n., sp. n., a fossil palm bug (Hemiptera: Thaumastocoridae: Xylastodorinae) in Dominican amber, with habits discernible by comparative functional morphology.  Entomologica Scandinavica 28:307-310.

47. Franqui, R. A., J. A. Santiago-Blay, S. Medina Gaud, and E. Abreu. 1997. The "Museo de Entomología y Biodiversidad Tropical" of the Agricultural Experiment Station, University of  Puerto Rico. Florida Entomologist 80:490-492. Web Site:

48. Virkki, N. and J. A. Santiago-Blay. 1997. Chromosome numbers of some Puerto Rican species of leaf beetles (Coleoptera; Chrysomelidae): Biogeographical and evolutionary considerations. Cytobios 90(361):103-131.

49. Virkki, N. and J. A. Santiago-Blay. 1998. Chromosome numbers in 71 Puerto Rican species of   leaf beetles (Coleoptera: Chrysomelidae). Journal of Agriculture of the University of Puerto Rico 82:69-83.

50. Lambert, J. B., C. E. Shawl, G. O. Poinar, Jr., and J. A. Santiago-Blay. 1999. Classification of modern resins by solid nuclear magnetic resonance spectroscopy. Bioorganic Chemistry 27:409-433.

51. Santiago-Blay, J. A., R. L. Hoffman, J. B. Lambert, and Y. Wu. 2003. “2002”. Cylindroiulus truncorum (Silvestri): a new milliped for Virginia (U.S.A.), with natural history observations (Julida: Julidae). Banisteria 20: 62-66.

52. Pohl, K., M. D. Therrell, J. [A.] Santiago-Blay, N. Ayotte, J. J. Cabrera Hernández, S. Díaz Castro, E. Cornejo Oviedo, J. A. Elvir, M. Gonzáles Elizondo, D. Opland, J. Park, G. Pederson, S. Bernal Salazar, L. Vazguez Selem, J. Villanueva Díaz, and D. Stahle. 2003. A cool season precipitation reconstruction for Saltillo, Mexico. 11th North American Dendroecological Fieldweek, Climatic Reconstruction Research Group, Mexico, August 2001. Tree-Ring Research 59(1):11-19.

53. Santiago-Blay, J. A., V. Fet, M. E. Soleglad, and P. R. Craig. 2004. The second Cretaceous scorpion specimen from Burmese amber (Arachnida: Scorpiones). Journal of Systematic Palaeontology 2(2):147-152.

54. Santiago-Blay, J. A., V. Fet, M. E. Soleglad, S. Anderson. 2004. A new genus and subfamily of scorpions from Cretaceous Burmese amber (Scorpiones: Chaerilidae). Revista Ibérica de Aracnología 9:3-14.

55. Cabrera-Asencio, I, J. A. Santiago-Blay, and A. L. Vélez. 2004 “2003”. Epitragus aurulentus (Kirsh) (Coleoptera: Tenebrionidae): A new record and report for Puerto Rico. Journal of Agriculture of the University of Puerto Rico 87(3-4):161-164.

56. Santiago-Blay, J. A., M. E. Soleglad, and V. Fet. 2004. A redescription and family placement of Uintascorpio  Perry, 1995 from the Parachute Creek Member of the Green River Formation (Middle Eocene) of Colorado, U.S.A. (Scorpiones: Buthidae). Revista Ibérica de Aracnología 10:7-16.

57. Lambert, J. B. Y. Wu, and J. A. Santiago-Blay. 2005. Taxonomic and chemical relationships revealed by nuclear magnetic resonance spectra of plant exudates.  Journal of Natural Products 68(5):635-648.

58. Santiago-Blay, J. A., S. R. Anderson, and R. T.  Buckley. 2006 “2005”. Possible implications of two new angiosperm flowers from Burmese amber (Lower Cretaceous) for well-established and diversified insect-plant interactions. Entomological News 116(5):341-346.

59. Baptista, C., J. A. Santiago-Blay, M. E. Soleglad, and V. Fet. 2006. The Cretaceous scorpion genus, Archaeobuthus,  revisited  (Scorpiones: Archaeobuthidae). Euscorpius 35:1-40.

60. Santiago-Blay, J. A.  2006. Marketing mostly intangible goods: the case of botanical gardens and arboreta.  Also, Case Teaching Notes for “Marketing Mostly Intangible Goods: The Case of Botanical Gardens and Arboreta”. Case Studies in Science. The National Center for Case Study Teaching in Science Case Collection. University of Buffalo. Buffalo, New York, U.S.A. Web versions can be found here, here, and here.

61. Lambert, J. B., M. A. Kozminski, C. A. Fahlstrom, and J. A. Santiago-Blay. 2007. Proton nuclear magnetic resonance characterization of resins from the family Pinaceae. Journal of Natural Products 70:188-195.

62. Lambert, J. B., Y. Wu, and M. A. Kozminski, and J. A. Santiago-Blay. 2007. Characterization of Eucalyptus and chemically related exudates by nuclear magnetic resonance spectroscopy. Australian Journal of Chemistry - an International Journal for Chemical Science 60:862-870.

63. Santiago-Blay, J. A. and J. B. Lambert. 2007. Amber’s botanical origins uncovered. American Scientist 95:150-157. Web versions can be found here . Reprinted, with permission as, Aux sources de l’ambre. Pour la Science [(Paris, France) French version of Scientific American] 356 (June 2007):70-75.A Abstracted by David M. Kondo in the winter 2007 issue of Gems and Gemology 43 (Winter 2007):395. (Not peer-reviewed publications.)

64. Santiago-Blay, J. A.  2008. Lubricating jelly helps improve image clarity of inclusions entombed in amber. Annales de la Société Entomologique de France (Nouvelle Série) 44(2):209-210.

65. Santiago-Blay, J. A. 2008. Aulacoscelinae Chapuis, 1874 (Insecta, Coleoptera, Orsodacnidae or Chrysomelidae): proposed conservation. Bulletin of Zoological Nomenclature65(2):97-105.

66. Lambert, J. B., J. A. Santiago-Blay, and K. B. Anderson. 2008. Chemical signatures of fossilized resins and recent plant exudates. Mini Review. Angewandte Chemie (International Edition) 47:9608-9616. Also published in German, with the following bibliographic information: Chemischer Fingerabdruck von fossilen Harzen und rezenten Pflanzenexsudaten. Angewandte Chemie 120:9750 – 9760

67. Santiago-Blay, J. A., B. C. Ratcliffe, F.-T. Krell, and R. Anderson. 2008. Allotypes should be from the type series: a position paper for reinstating Recommendation 72A from the third edition of the Code that defines the term ‘allotype’. Bulletin of Zoological Nomenclature65(4):260-264.

68. McCafferty, W. P. and J. A. Santiago-Blay. 2009. A new Cretaceous mayfly fossil from Burmese amber (Ephemeroptera: Australiphemeridae). Entomological News 119(5):492-496.

69. Santiago-Blay, J. A.  2009. Systematics and some aspects of the biology of the scorpions (Arachnida) of the Greater Puerto Rico Region: A biosystematic synopsis. Entomological News 120(1):109-124, and back cover. Erratum: Entomological News 120(1):152.

70. Lambert, J. B., E. R. Heckenbach A. E. Hurtley, Y. Wu, and J. A. Santiago-Blay. 2009. Nuclear magnetic resonance spectroscopic characterization of legume exudates. Journal of Natural Products 72:1028-1035.

71. Ren, D., C. C. Labandeira, J. A. Santiago-Blay, A. Rasnitsyn, C-K. Shih, A. S. Bashkuev, M. A. V. Logan, C. L. Hotton, and D. Dilcher. 2009. A probable pollination mode before angiosperms: Eurasian, long-proboscid scorpionflies. Science 326:840-847 and supplemental material. A description was published in the November 5, 2009 web edition of Science News written by S. Perkins