Abstract

The beetle genus Pharaxonotha Reitter (Coleoptera: Erotylidae: Pharaxonothinae) is found in the cones of cycad genera in the New World, including species of Ceratozamia Brongn., Dioon Lindl., Microcy¬cas (Miq.) A.DC and Zamia L. Its presence and diversity are analyzed for the 17 species of Zamia known to occur in Panama. Nine species are recognized, and five new species are described: Pharaxonotha clarkorum Pakaluk, P. confusa Pakaluk, P. fortunensis Tang, Skelley and Taylor new species, P. holzmani Tang, Skelley and Taylor new species, P. kirschii Reitter, P. manicatae Tang, Skelley and Taylor new species, P. panamensis Tang, Skelley and Taylor new species, P. pseudoparasitica Tang, Skelley and Taylor new species, and P. taylori Skelley and Tang. A key to species of Pharaxonotha in Panama is presented.

References

• Acuña-Castillo RH, Gómez LD. 2009. Selección de un lectotipo para Zamia fairchildiana L.D. Gómez (Zamiaceae). Brenesia 71–72: 65-66.  
• Anderson DM. 1987. Chapter 4. Larval beetles (Coleoptera). p. 95–113. In: Gorham JR (ed.). Insect and mite pests in food: An illustrated key. U.S. Department of Agriculture. Agriculture Handbook 655: 1–767.  
• Basset Y, Aberlene HP, Delvare G. 1992. Abundance and stratification of foliage arthropods in a lowland rain forest of Cameroon. Ecological Entomology 17: 310–318.  
• Booth RG, Cox ML, Madge RB. 1990. IIE guides to insects of importance to man. 3. Coleoptera. International Institute of Entomology; The Natural History Museum; London. 384 p 
• Calonje M, Meerow AW, Griffith MP, Salas-Leiva D, Vovides AP, Coiro M, Francisco-Ortega J. 2019. A time-calibrated species tree phylogeny of the New World cycad genus Zamia L. (Zamiaceae, Cycadales). International Journal of Plant Sciences 180: 286–314.  
• Calonje M, Stevenson D, Calonje C, Ramos, YA, Lindstrom A. 2010. A new species of Zamia from Chocó, Colombia (Cycadales, Zamiaceae). Brittonia 62: 80–85.  
• Calonje M, Stevenson D, Osborne R. 2023. The World List of Cycads. Available at https://www.cycadlist.org/  (Last accessed November 30, 2023.)
• Champion GC. 1904. Pharaxonotha kirschii, Reitt. p. 36 [footnote]. In: Arrow GJ. Note on two species of Coleoptera introduced into Europe. Entomologist’s Monthly Magazine 40: 35–36.  
• Chaves R, Genaro JA. 2005. A new species of Pharaxonotha (Coleoptera: Erotylidae), probable pollinator of the endangered Cuban cycad, Microcycas calocoma (Zamiaceae). Insecta Mundi 19(3): 143–150.  
• Chemnick J, Hoese H. 2008. Sympatry in the Cycadales. Cycad Newsletter 31(2/3): 26–27.  
• Chittenden FH. 1895. On the distribution of certain imported beetles. Insect Life VII(4): 326–332.  
• Chittenden FH. 1911. The Mexican grain beetle (Pharaxonotha kirschi Reitt.). Bulletin of the United States Department of Agriculture. Bureau of Entomology, Washington, DC. N. ser. 96(1): 8–13.  
• Coates A. 1997. The forging of Central America. p. 1–37. In: Coates A (ed.). Central America: a natural and cultural history. Yale University Press; New Haven, CT. 277 p 
• Cruz-Angón A, Baena ML, Greenberg R. 2009. The contribution of epiphytes to the abundance and species richness of canopy insects in a Mexican coffee plantation. Journal of Tropical Ecology 25: 453–463.  
• Franz NM, Skelley PE. 2008. Pharaxonotha portophylla (Coleoptera: Erotylidae), new species and pollinator of Zamia (Zamiaceae) in Puerto Rico. Caribbean Journal of Science 44(3): 321–333.  
• Gómez LD. 1982. Plantae Mesoamericanae Novae. II. Phytologia 50(6): 401–404.  
• Gómez LD. 1986. Zamia pseudomonticola. p. 141. In: Stevenson DW, Sabato S. Typification of names in Zamia L. and Aulacophyllum Regel (Zamiaceae). Taxon 35(1): 134–144.  
• Gorham HS. 1898. Coccinellidae (part) and supplement to Erotylidae (part). p. 241–256. In: Godman F, Salvin O (eds.). Biologia Centrali-Americana, Insecta, Coleoptera, vol. 7. R. H. Porter; London. 276 p. + 13 pl 
• Griffith MP, Calonje MA, Stevenson DW, Husby CE, Little DP. 2012. Time, place, and relationships: cycad phenology in a phylogenetic and biogeographic context. Memoirs of the New York Botanical Garden 106: 59–81.  
• Hanley RS, Ashe JS. 2003. Techniques for dissecting adult aleocharine beetles (Coleoptera: Staphylinidae). Bulletin of Entomological Research 93: 11–18.  
• Hinton HE. 1945. A monograph of the beetles associated with stored products. British Museum (Natural History); London. 442 p 
• Kingsolver JM. 1973. New synonymy in Languriidae (Coleoptera). Proceedings of the Entomological Society of Washington 75(2): 247.  
• Lawrence JF. 1991. Languriidae (Coleoptera) (including Cryptophilidae). p. 471–473. In: Stehr FW (ed.). Immature insects. Vol. 2. Kendall/Hunt Publishing Company; Dubuque, IA. XVI + 975 p 
• Lawrence JF, Beutel RG, Leschen RAB, Ślipiński SA. 2010. Chapter 2. Glossary of morphological terms. p. 9–20. In: Leschen RAB, Beutel RG, Lawrence JF (eds.). Handbook of zoology, Coleoptera volume 2: morphology and systematics (Elateroidea, Bostrichiformia, Cucujiformia partim). Walter de Gruyter; Berlin, Germany. 785 p 
• McHugh JV, Marshall CJ, Fawcett FL. 1997. A study of the adult morphology in Megalodacne heros (Say) (Coleoptera: Erotylidae). Transactions of the American Entomological Society 123: 167–223.  
• McKenna DD, Shin S, Ahrens D, Balke M, Beza-Beza C, Clarke DJ, Donath A, Escalona HE, Friedrich F, Letsch H, Liu S, Maddison D, Mayer C, Misof B, Murin PJ, Niehuis O, Peters RS, Podsiadlowski L, Pohl LH, Scully ED, Yan EV, Zhou X, Ślipiński A, Beutel RG. 2019. The evolution and genomic basis of beetle diversity. Proceedings of the National Academy of Sciences 116: 24729–24737.  
• Merckx VSFT, Hendriks KP, Beentjes KK, Mennes CB, Becking LE, Peijnenburg KTCA, Afendy A, Arumugam N, de Boer H, Biun A, Buang MM, Chen PP, Chung AYC, Dow R, Feijen FAA, Feijen H, Feijen-van Soest C, Geml J, Geurts R, Gravendeel B, Hovenkamp P, Imbun P, Ipor I, Janssens SB, Jocque´ M, Kappes H, Khoo E, Koomen P, Lens F, Majapun RJ, Morgado LN, Neupane S, Nieser N, Pereira JT, Rahman H, Sabran S, Sawang A, Schwallier RM, Shim PS, Smit H, Sol N, Spait M, Stech M, Stokvis F, Sugau JB, Suleiman M, Sumail S, Thomas DC, van Tol J, Tuh FYY, Yahya BE, Nais J, Repin R, Lakim M, Schilthuizen M. 2015. Evolution of endemism on a young tropical mountain. Nature 524: 347–350.  
• Norstog K, Fawcett PKS. 1989. Insect-cycad symbiosis and its relation to the pollination of Zamia furfuracea (Zamiaceae) by Rhopalotria mollis (Curculionidae). American Journal of Botany 76(9): 1380–1394.  
• Norstog K, Fawcett PKS, Vovides AP. 1992. Beetle pollination of two species of Zamia: Evolutionary and ecological considerations. The Palaeobotanist 41: 149–158.  
• Norstog K, Nichols TJ. 1997. The biology of the cycads. Cornell University Press; Ithaca, NY. 363 p 
• O’Brien C, Tang W. 2015. Revision of the New World cycad weevils of the subtribe Allocorynina, with description of two new genera and three new subgenera (Coleoptera: Belidae: Oxycoryninae). Zootaxa 3970: 1–87.  
• Pakaluk J. 1988. Review of the New World species of Pharaxonotha Reitter (Coleoptera: Languriidae). Revista de Biologia Tropical 36(2B): 447–451.  
• Pindell J, Kennan L. 2009. Tectonic evolution of the Gulf of Mexico, Caribbean and northern South America in the mantle reference frame: an update. p. 1–55. In: James K, Lorente MA, Pindell J (eds.). The Origin and Evolution of the Caribbean plate. Geological Society of London, Special Publication 328: 1–868.  
• Powell GS. 2021. Phylogeny and visual system evolution in sap beetles (Coleoptera: Cucujoidea: Nitidulidae). Doctoral dissertation, Brigham Young University; Provo, Utah, USA. 175 p. Available at https://scholarsarchive.byu.edu/cgi/viewcontent.cgi?article=10823&context=etd.  (Last accessed October 28, 2023.)
• Reitter E. 1875. Revision der europäischen Cryptophagiden. Deutsche Entomologische Zeitschrift 19(3): 1–86.  
• Segalla R, Pinheiro F, Morellato LPC. 2021. Reproductive biology of the South American cycad Zamia boliviana, involving brood-site pollination. Plant Species Biology 36: 348–360.  
• Sharp, D. 1900. Cryptophagidae. p. 579–624, pl. 18. In: Godman F, Salvin O (eds.). Biologia Centrali-Americana. Insecta. Coleoptera. Vol II. Part 1 [1887–1905]. R. H. Porter; London. 717 p. + 19 pl. [<i>Pharaxonotha </i>p. 598] 
• Skelley PE, Segalla R. 2019. A new species of Pharaxonotha Reitter (Coleoptera: Erotylidae) from central South America. Zootaxa 4590(1): 184–190.  
• Skelley PE, Tang W. 2020. Two new species of Pharaxonotha Reitter among the early-diverging lineages, with a key to the species of the genus (Coleoptera: Erotylidae: Pharaxonothinae). Insecta Mundi 0837: 1–11.  
• Skelley PE, Tang W, Pérez-Farrera MA. 2022. Review of Pharaxonotha Reitter (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting the cycad genus Dioon Lindl. (Cycadales), with descriptions of nine new species and comments on P. kirschii Reitter. Insecta Mundi 0917: 1–41.  
• Skelley PE, Xu G, Tang W, Lindström A, Marler T, Khuraijam JS, Singh R, Radha P, Rich S. 2017. Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267: 1–63.  
• Stevenson DW. 1993. The Zamiaceae in Panama with comments on phytogeography and species relationships. Brittonia 45: 1–16.  
• Sutton SL, Ash CPJ, Grundy A. 1983. The vertical distribution of flying insects in lowland rain-forests of Panama, Papua New Guinea and Brunei. Zoological Journal of the Linnean Society 78: 287–297.  
• Tang W. 1987. Insect pollination in the cycad Zamia pumila (Zamiaceae). American Journal of Botany 74: 90–99.  
• Tang W, Gomez Dominguez H. 2022. Two new species of Pharaxonothinae beetles (Coleoptera: Erotylidae) inhabiting cones of the cycad Ceratozamia santillanii Perez-Far. & Vovides (Cycadales: Zamiaceae) in Mexico. Zootaxa 5150(3): 428–442.  
• Tang W, Skelley P, Pérez-Farrera MA. 2018a. Ceratophila, a new genus of erotylid beetles (Erotylidae: Pharaxonothinae) inhabiting male cones of the cycad Ceratozamia (Cycadales: Zamiaceae). Zootaxa 4508: 151–178.  
• Tang W, Xu G, O’Brien CW, Calonje M, Franz NM, Johnston MA, Taylor A, Vovides AP, Pérez-Farrera MA, Salas- Morales SH, Lazcano-Lara JC, Skelley P, Lopez-Gallego C, Lindström A, Rich S. 2018b. Molecular and morphological phylogenetic analyses of New World cycad beetles: what they reveal about cycad evolution in the New World. Diversity 10(38): 1–26.  
• Tang W, Xu W, Marler T, Khuraijam JS, Singh R, Lindström AJ, Radha P, Rich S, Nguyen KS, Skelley P. 2020. Beetles (Coleoptera) in cones of cycads (Cycadales) of the northern hemisphere: diversity and evolution. Insecta Mundi 0781: 1–19.  
• Taylor AS, Calonje M. 2015. The meaning of artificial hybridization in cycad phylogenetic studies: The case for populations of Zamia cunaria and Zamia ipetiensis. p. 47. In: Calonje M (ed.). Cycad 2015 10th international conference on cycad biology schedule and abstracts. Montgomery Botanical Center; Coral Gables, FL. 87 p 
• Taylor AS, Haynes JL, Holzman G. 2008. Taxonomical, nomenclatural and biogeographical revelations in the Zamia skinneri complex of Central America (Cycadales: Zamiaceae). Botanical Journal of the Linnean Society 158: 399–429.  
• Taylor AS, Holzman G. 2012. A new Zamia species from the Panama Canal Area. Botanical Review 78: 335–344.  
• Terry I, Tang W, Taylor Blake A, Donaldson JS, Singh R, Vovides AP, Cibrián Jaramillo A. 2012. An overview of cycad pollination studies. p. 352–394. In: Stevenson DW, R. Osborne R, Taylor Blake AS (eds.). Proceedings of the 8th International Conference on Cycad Biology. Memoirs of The New York Botanical Garden, 106. The New York Botanical Garden Press; New York, NY. 554 p 
• Valencia-Montoya WA, Tuberquia D, Guzmán PA, Cardona-Duque J. 2017. Pollination of the cycad Zamia incognita A. Lindstr. & Idárraga by Pharaxonotha beetles in the Magdalena Medio Valley, Colombia: a mutualism dependent on a specific pollinator and its significance for conservation. Arthropod-Plant Interactions 5: 717–729.  
• Wheeler QD, Platnick NI. 2000. The phylogenetic species concept (sensu Wheeler and Platnick). p. 55–69. In: Wheeler QD, Meier R (eds.). Species concepts and phylogenetic theory: a debate. Columbia University Press; New York, NY. 256 p 
• Wolda H, O’Brien CW, Stockwell HP. 1998. Weevil diversity and seasonality in tropical Panama as deduced from light-trap catches (Coleoptera: Curculionoidea). Smithsonian Contributions to Zoology 590: 1–79.  
• Xu G, Tang W, Skelley P, Liu N, Rich S. 2015. Cycadophila, a new genus (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas debaoensis (Cycadaceae) in Asia. Zootaxa 3986: 251–278.