On the diversity of Neotropical Hymenoptera

Fernández, Fernando; Fernández, Fernando

doi:10.15446/caldasia.v44n3.94286

Serviços Personalizados

Journal

Artigo

Indicadores

Citado por SciELO
Acessos

Links relacionados

Citado por Google
Similares em SciELO
Similares em Google

Mais
Mais

Permalink

Caldasia

versão impressa ISSN 0366-5232versão On-line ISSN 2357-3759

Caldasia vol.44 no.3 Bogotá set./dez. 2022 Epub 06-Jun-2024

https://doi.org/10.15446/caldasia.v44n3.94286

Taxonomy and systematics

On the diversity of Neotropical Hymenoptera

Sobre la diversidad de Hymenoptera neotropicales

Fernando Fernández¹
http://orcid.org/0000-0002-7581-1428

^¹Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá D.C., Colombia, ffernandezca@unal.edu.co

ABSTRACT

Hymenoptera is globally one of the most diverse groups of insects. There are no recent estimates nor sound data on the size of the fauna of this order in the Neotropical Region. Based on printed or digital catalogs, lists, as well as reviews of monographs, and other sources, the Neotropics comprises 26 su-perfamilies, 92 families, 3162 genera, and over 33 640 species of Hymenoptera. Hymenoptera diversity in the Neotropics is greater than in the Nearctic, Palearctic and Australian regions. The Neotropics is probably the richest region in the World in families, genera, and species of the order, with the majority of taxa endemic to this region.

Keywords: Diversity; Hymenoptera; Neotropics; taxonomy

RESUMEN

Hymenoptera es globalmente uno de los grupos más diversos de insectos. No hay estimaciones o datos firmes recientes sobre el tamaño de la fauna de este orden en la Región Neotropical. Tomando como base catálogos, listas impresas o en forma digital, además de monografías y otras fuentes, el Neotrópico comprende 26 superfamilias, 92 familias, 3162 géneros y un poco más de 33 640 especies. La diversidad de Hymenoptera en el Neotrópico es mayor que en las regiones Neártica, Paleártica y Australiana. Probablemente es el Neotrópico la región más rica del mundo en familias, géneros y especies de Hymenoptera, con la mayoría de taxones endémicos de esta región.

Palabras clave: Hymenoptera; Neotrópico; diversidad; taxonomía

Wasps, bees and ants, order Hymenoptera, are one of the most diverse and important groups of insects in terrestrial ecosystems. The vast majority of species are parasitoid wasps that exert a natural control over populations of other insects, many of them phytophagous. Bees are important pollinators of many plants. Ants are key players in the structure and dynamics of forests. Their wealth of behavioral patterns (solitary, communal, subsocial, social, and parasitic) is an obligatory source for the study of the evolution of sociability. In addition, some species are agricultural pests, or may cause veterinary or medical problems. In contrast to the importance of the order, knowledge of its diversity at the global and regional level is key. Knowing how many species there are, what names they have, and where they are found is vital for studies in systematics, phylogeny, biogeography, ecology and conservation.

As Hymenoptera is one of the four hyper-diverse insect orders, it is not easy to establish precise numbers of the described species in the group. It is even more challenging to have estimates of undescribed species, a task that has become increasingly difficult in face of the decline of biological collections and curators, mainly taxonomists. In Hymenoptera, 111 families, 8423 genera (^{Aguiar et al. 2013}) and 154 000 living species (^{Huber 2017}) have been described, making it the most diverse order after Coleoptera and Lepidoptera, and close to Diptera. Estimates of total species richness are also variable, generally exceeding one million (^{Ulrich 1999}). One recent study suggests a hidden high diversity of parasitoid wasps (^{Forbes et al. 2018}). If true, this planet is flooded not with beetles, but with wasps.

For Coleoptera there is at least one checklist to the Neotropical Region published by ^{Blackwelder between 1944} and 1957 and reprinted by the Smithsonian. In Lepidoptera there are innumerable catalogs with an emphasis on butterflies, being J.B. Heppner the editor of a series that covers most superfamilies and families of the order, with the exception of Geometroidea and Noctuoidea which are covered in other sources (e.g. ^{Scobble 1999} for Geometridae). Nelson Papavero during several decades published the serial catalog of Diptera of the Neotropical region (^{Klassa and dos Santos 2014}).

For Neotropical Hymenoptera there are printed or digital catalogs for specific groups such as sawflies (^{Taeger et al. 2010}), ants (^{Kempf 1972}, ^{Bolton 1995}, c2022, ^{Bolton et al. 2007}), bees (^{Moure et al. 2007}) and non-bee apoids (^{Amarante 2002}, 2005), or online resources, such as for Chalcidoidea (^{Noyes c2020}). But there are no printed or digital resources that list the entire fauna of Hymenoptera described for the Neotropics. How many species of Hymenoptera have been described for the Neotropics? ^{Fernández (2000)} offers a rough assessment, indicating 20 super families, 77 families, 2527 genera and around 24 000 species, estimating the number of species for the region at 60 000. These numbers are now out of date due to the many taxonomic developments since then, but also due to better access to publications and databases with information on poorly known groups at that time (such as Ceraphronoidea) or very rich groups with old and inaccessible literature (such as Chalcidoidea).

A new assessment of the size of Neotropical Hymenoptera fauna has improved from access to more sources of information, either in old checklists now available online, new lists, or the extraction of information from online catalogs, such as those dealing with bees and chalcidoids. Here is offered the list of Hymenoptera superfam-ilies and families known in the Neotropical region with the number of genera and described species with the sources before each family (Table 1). In most cases, the numbers are taken from the original references. In most taxa, the data from each list was verified for country or, as in the case of ^{Johnson (1992)}, for region. Here we report the number of families, genera and species for each superfamily (Table 1). From the Noyes Chalcidoidea database, the names for each family, genus and species were consulted under the "Neotropical" restriction, and lists were compiled for each family. This procedure was followed for the online catalog of bees of the Neotropical Region, which was last updated in 2013.

According to table 1 there are 26 superfamilies, 92 families, 3162 genera and 33645 species for the Neotropics (see also supplementary Figs. S1-4). These figures imply a notable increase regarding the numbers presented in ^{Fernández (2000)}. Part of these differences are explained by new studies in phylogeny that have created new suprageneric categories (^{Pilgrim et al. 2008}, ^{Sann et al. 2018}, ^{Chen et al. 2021}, ^{Zhang et al. 2022}).

The number of described genera and species can be considered reliable for groups in which there is recent, revised and constantly updated information, such as the Chalcidoidea database of Noyes or ^{Bolton's catalog of ants (Bolton c2022)}.

Table 1 Superfamilies and families of Neotropical Hymenoptera, with number of genera, species and references.

However, in key groups such as bees or microhymenoptera (excluding Chalcidoidea) there is no updated information. In some cases (such as Diaprioidea and Platygastroidea), efforts have been made to scan recent literature. In the case of Cynipoidea, one of the important groups of unlisted wasps, the work of making a checklist based on original literature is being undertaken. There are a few families of Hymenoptera for which there are no catalogs or lists available, such as Tiphiidae or Thynnidae.

In the megadiverse superfamily Ichneumonoidea, the task of establishing updated numbers of genera, and especially of species, is difficult. Making estimates of diversity is even more complicated, due to the apparent large number of species that remain undescribed in the tropics (e.g. ^{Zaldívar-Riverón et al. 2010}, Fernández-Florez et al. 2013, ^{Marsh et al. 2013}, ^{Sharkey et al. 2021}). However, the high numbers of new species reported with molecular techniques, as in the case of the 403 new species of Braconidae described in ^{Sharkey et al. (2021)} using COI barcode clusters should be viewed carefully. ^{Meier et al. (2021)} show that many of Sharkey et al.'s species are unstable when the underlying data are analyzed using different species delimitation algorithms.

This creates a new problem, that of the proliferation of new names weakly supported (the "superficial description impediment": ^{Meier et al. 2021}).

The most species-rich families are Ichneumonidae (4505), Braconidae (4142), Formicidae (3200), Crabronidae (1304), Apidae (1926), Eulophidae (1541), Mutillidae (1505), Vespidae (1427), Encyrtidae (1200), Megachilidae (1014), Halictidae (1004) and Pompilidae (nearly 1000) (Table 1, Supplementary Fig. S1). Regarding the number of genera, the richest families are Ichneumonidae (472), Braconidae (465), Pteromalidae (230), Encyrtidae (170), Formicidae (142), Apidae (135), and Eulophidae (120) (Table 1, Supplementary Fig. S2). There are families rich in species, but not in genera, such as Crabronidae, or rich in genera and not in species, such as Pteromalidae. Others occupy the first places in any criterion of number of genera or species, such as Ichneumonidae and Braconidae, groups considered hyper-diverse.

Despite the limitations noted above, and the certain lack of recent information on some groups (such as Tiphiidae), the numbers offered here are a good estimate of the described richness of the order Hymenoptera. When making adjustments (new taxa vs synonymized taxa), the number of described genera should be close to 3200 and the number of described species close to 34 000. These numbers respectively represent 38 % of World genera and 22 % World species. Even though there is no recent and comparable information on the diversity of Hymenoptera in each of the large natural regions, the following data gives an idea of the sizes of their fauna: About 13 000 species in Australia (8 % of the World, ^{Britton c2018}), 17 429 in the Nearctic (12 % of the World, ^{Danks and Smith 2009}), just over 21 000 in the Palearctic (14 % of the World, ^{Korotyaev et al. 2017}). Even without knowing the numbers for the other two great regions, the Afrotropical and Oriental, it is highly probable their combined Hymenoptera fauna is the richest in the world, with almost 34 000 described species.

Another matter is estimating the total number of species in the Neotropics and other regions. For the Nearctic, ^{Grissell (1999)} estimates the number of species may be about 36 000 (50 % more). This author cites 17 000 described species only for Costa Rica and estimates total number between 20 000 and 40 000. If these numbers and approximations are accurate, the Neotropical fauna could easily reach 100 000 species, more than 60 % of which is not described. Since global estimates of the richness of Hymenoptera is highly variable (between 30 000 and 2.5 million species) the estimated number for the Neotropics can vary between 100 000 and almost a million. In the most recent approach ^{Forbes et al. (2018)} extrapolate information on the diversity of parasitoids and estimates 1 152 127 species of Hymenoptera for the World.

It is striking that the Neotropics are home to a rich fauna, with a high number of endemism, but at the same time it is facing serious problems that threaten the future of biological diversity (^{Mundy 2020}). Deforestation, change in land management, illegal and legal mining, climate change, introduction of invasive species, new pathogens, overpopulation and pollution are exerting strong damaging pressures on the diversity of the region, from humid lowland forests to páramos (e.g. ^{Alroy 2017}). The disappearance of Hymenoptera species, as well as other insects (^{Wagner et al. 2021}), not only compromises the stability of ecosystems, but also ecosystem services of interest to humans, such as the natural control of forest pests or the pollination of plants of economic interest.

ACKNOWLEDGEMENTS

To colleagues Edgar Palacio, Diego Campos, Carlos Sarmiento, and Jim Carpenter for providing updated data on the families Ichneumonidae, Braconidae and Vespidae. To Lubo Masner, Dave Smith, Mike Sharkey, Norman Johnson, José Luis Nieves-Aldrey, and P. Buhl for their support in obtaining literature in various Hymenoptera groups. To Klaus Rassmusen and an anonymous reviewer for improving the manuscript. To Andrés Sánchez for their kindly elaboration of the figures.

LITERATURE CITED

Aguiar AP, Deans AR, Engel MS, Forshage M, Huber JT, Jennings JT, Johnson NF, Lelej AS, Longino JT, Lohrmann V, Mikó I, Ohl M, Rasmussen C, Taeger A, Yu DSK. 2013. Order Hymenoptera. In: Zhang Z-Q, editor. Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness (Addenda 2013). Zootaxa 3703(1):51-62. doi: https://doi.org/10.11646/zootaxa.3703.L12 [ Links ]

Aguiar AP. 1998. Revisão do gênero Hemistephanus Enderlein, 1906 (Hymenoptera: Stephanidae), com considerações metodológicas. Rev. Bras. Entomol. 41(2-4):343-429. [ Links ]

Aguiar AP. 2004. World catalog of the Stephanidae (Hymenop-tera: Stephanoidea). Zootaxa 753:1-120. doi: https://doi.org/10.11646/zootaxa.753.1.1 [ Links ]

Allen WH. 1972. A monographic study of the subfamily Tiphiinae (Hymenoptera: Tiphiidae) of South America. Smith-son. Contrib. Zool. 113:1-76. doi: https://doi.org/10.5479/si.00810282.113 [ Links ]

Alroy J. 2017. Effects of habitat disturbance on tropical forest biodiversity. PNAS 114(23):6056-6061. doi: https://doi.org/10.1073/pnas.1611855114 [ Links ]

Amarante STP. 2002. A synonymic catalog of the Neotropical Crabronidae and Sphecidae. (Hymenoptera: Apoidea). Arq. Zool. 37(1):1-139. doi: https://doi.org/10.11606/issn.2176-7793.v37i1p1-139 [ Links ]

Amarante STP. 2005. Addendum and corrections to a synonymic catalog of Neotropical Crabronidae and Sphecidae. Pap. Avulsos Zool. (São Paulo) 45(1):1-18. doi: https://doi.org/10.1590/S0031-10492005000100001 [ Links ]

Argaman Q. 1988. Generic synopsis of Sclerogibbidae (Hymenoptera). Ann. Hist. Nat. Mus. Natl. Hung. 80:177-187. [ Links ]

Arias D. 2002. Lista de los géneros y especies de la superfamilia Platygastroidea (Hymenoptera) de la Región Neotropical. Biot. Colomb. 3(2):215-233. [ Links ]

Arias DC, Delvare G. 2003. List of the genera and species of family Chalcididae (Hymenoptera: Chalcidoidea) of the Neotropical region. Biot. Colomb. 4(2):123-144. [ Links ]

Azevedo CO, Molin AD, Penteado-Dias A, Macedo ACC, Rodriguez-V B, Dias BZK, Waichert C, Aquino D, Smith DR, Shimbori EM, FB Noll, Gibson G, Onody HC, Carpenter JM, Lattke JE, Ramos KdosS, Williams K, Masner L, Kimsey LS, Tavares T, Olmi M, Buffington ML, Ohl M, Sharkey M, Johnson NF, Kawada R, Gonçalves RB, Feitosa RM, Heydon S, Guerra TM, Silva TS.R. da, Costa V. 2015. Checklist of the genera of Hymenoptera (Insecta) from Espírito Santo state, Brazil. Bol. Mus. Biol. Mello leitão (N. sér.) 37(3):313-343. [ Links ]

Azevedo CO. 1999. A key to world species of Scolebythidae (Hymenoptera: Chrysidoidea), with description of a new species of Dominibythus from Brazil. J. Hymenopt. Res. 8:1-5. [ Links ]

Blackwelder RE. 1944-1957. Checklist of the coleopterous insects of Mexico, Central America, the West Indies, and South America. Parts 1 through 6. Bull. U. S. Nat. Mus. 927-1492. doi: https://doi.org/10.5479/si.03629236.185.927 [ Links ]

Bohart R, Menke AS. 1976. Sphecids wasps of the world. A generic revision. California: University of California Press. [ Links ]

Bolton B, Alpert G, Ward PS, Naskrecki P. 2007. Bolton's Catalogue of Ants of the World. Cambridge-London: Harvard University Press, Cambridge-London, CD-Rom. [ Links ]

Bolton B. 1995. A new general catalogue of the ants of the world. Cambridge-London: Cambridge-London, Harvard University Press. [ Links ]

Bolton B. c2022. An online catalog of the ants of the world. [Last accessed: 27 Jan 2022] [Last accessed: 27 Jan 2022] https://www.antcat.org/ [ Links ]

Bouček Z, Noyes JS. 1987. Rotoitidae, a curious new family Chalcidoidea (Hymenoptera) from New Zealand. Syst. Entomol. 12:407-412. doi: https://doi.org/10.1m/j.1365-3113.1987.tb00212.x [ Links ]

Bouček Z. 1974. A revision of the Leucospidae (Hymenoptera: Chalcidoidea). Bol. Mus. Biol. Mello Suppl. 23:1-241. [ Links ]

Bouček, Z. 1975. The pteromalid subfamily Eutrichosomatinae (Hymenoptera: Chalcidoidea). J. Ent. 43:129-138. [ Links ]

Bradley JC. 1945. The Scoliidae (Hymenoptera) of Northern South America, with special reference to Venezuela. I. The genus Campsomeris. Bol. Entomol. Venez. 4(1):1-36. [ Links ]

Britton D. c2018. Ants, wasps, bees and sawflies: Order Hymenoptera. [Last accessed: 28 Jan 2021]. https://australian.museum/learn/animals/insects/sawflies-wasps-bees-ants-hymenoptera/ [ Links ]

Brothers DJ, Lelej AS. 2017. Phylogeny and higher classification of Mutillidae (Hymenoptera) based on morphological reanalyses. J. Hymenopt. Res. 60:1-97. doi: https://doi.org/10.3897/jhr.60.20091 [ Links ]

Brothers DJ. 1970. Discovery of the mutillid subfamily Typhoctinae in South America, with description of a new species (Hymenoptera). J. Kansas. Entomol. Soc. 43:302-308. [ Links ]

Buffington ML, Forshage M, Liljeblad J, Tang C-T, van Noort S. 2020. World Cynipoidea (Hymenoptera): A key to higher-level groups. Insect Syst. Divers. 4(4)1:1-69. doi: https://doi.org/10.1093/isd/ixaa003 [ Links ]

Buhl PN. 2011. New Neotropical species of Platygastrinae and Sce-liotrachelinae (Hymenoptera: Platygastridae), with keys to species of the larger genera, some redescriptions and a checklist. Folia Heyrovskyana (A) 19(1-4):25-128. [ Links ]

Carmean D, Kimsey L. 1998. Phylogenetic revision of the parasitoid wasp family Trigonalidae (Hymenoptera). Syst. Entomol. 23 (1): 35-76. doi: https://doi.org/10.1046/j.1365-3113.1998.00042.x [ Links ]

Chen H, Lahey Z, Talamas EJ, Valerio AA, Popovici OA, Musetti L, Klompen H, Polaszek A, Masner L, Austin AD, Johnson NF. 2021. An integrated phylogenetic reassessment of the parasitoid superfamily Platygastroidea (Hymenoptera: Proctotrupomorpha) results in a revised familial classification. Syst. Entomol. 46:1088-1113. doi: https://doi.org/10.1111/syen.12511 [ Links ]

Danforth BN, Minckley RL, Neff JL. 2019. The solitary bees: Biology, evolution, conservation. Princeton, EE. UU: Princeton University Press, Princeton. [ Links ]

Danks HV, Smith ABT. 2009. Insect biodiversity in the Nearctic Region. Chapter 3. In: Foottit RG, Adler PH, editors. Insect biodiversity: science and society, Second Edition. Wiley. Estados Unidos: John Wiley & Sons. p. 35-48. [ Links ]

De Santis L. 1979. Catálogo de los himénopteros calcidoideos de América al sur de los Estados Unidos. Buenos Aires, Argentina: Publicación Especial Comisión de Investigaciones Científicas Provincia de Buenos Aires, Argentina. [ Links ]

De Santis L. 1983. Catálogo de los himenópteros Calcidoideos de America al Sur de los Estados Unidos - Primer Suplemento. Rev. Per. Entomol. 24(1):1-38. [ Links ]

De Santis L. 1989. Catálogo de los himenópteros Calcidoides (Hymenoptera) al sur de los Estados Unidos, segundo suplemento. Catalogue of the Chalcidoidea (Hymenoptera) of America south of the United States, second supplement. Acta Entomol. Chil. 15:9-89. [ Links ]

Deans AR, Hueben M. 2003. Annotated key to the ensign wasp (Hymenoptera: Evaniidae) genera of the world, with descriptions of three new genera. P. Entomol. Soc. Wash. 105: 859-875. [ Links ]

Deans AR. 2005. Annotated catalog of the world's ensign wasp species (Hymenoptera: Evanioidea). Am. Entom. Inst. 34:1-158. [ Links ]

Delvare G, Boucek Z. 1992. On the New World Chalcididae. Mem. Am. Entomol. Inst. 53:1-466. [ Links ]

Diaz NB, Gallardo FE, Durante SP. 2002. Estado del conocimiento de los Cynipoidea en la Región Neotropical (Hymenoptera). España: Proyecto de Red Iberoamericana de Biogeografía y Entomología Sistemática PrIBES. [ Links ]

Diez P, Fidalgo P, Roig-Alsina A. 2007. A new genus and species of Plumariidae (Hymenoptera, Chrysidoidea) from western xeric Argentina. Zootaxa 1467:35-41. doi: https://doi.org/10.11646/zootaxa.1467.1.3 [ Links ]

Evans HE. 1961. A preliminary review of the Nearctic species of Sierolomorpha (Hymenoptera). Breviora 140:1-12. [ Links ]

Evans HE. 1963. A new family of wasps. Psyche 70:7-16. doi: https://doi.org/10.1155/1963/92486 [ Links ]

Evans HE. 1964. A synopsis of the American Bethylidae (Hyme-noptera, Aculeata). Bull. Mus. Comp. Zool. 132(1):1-222. [ Links ]

Fernández F, Guerrero RJ, Sánchez-Restrepo A. 2021. Systematics and diversity of Neotropical ants. Rev. Col. Ent. 47(1):e11082. doi: https://doi.org/10.25100/socolen.v47i1.11082 [ Links ]

Fernández F, Rodríguez J, Waichert C, Decker B, Pitts J. 2022. Twenty two years later: An updated checklist of Neotropical spider wasps (Hymenoptera: Pompilidae). Zootaxa 5116(4):451-503. [ Links ]

Fernández F, Sarmiento CE. 2002. A new species of Olixon Cameron (Hymenoptera: Rhopalosomatidae) from the Neotropical Region. J. N.Y. Entomol. Soc. 110(3-4):413-416. [ Links ]

Fernández F, Sarmiento CE. 2015. A new species of Huarpea Pate, 1947 from Colombia and redescription of H. wagneriella (du Buysson, 1904) (Hymenoptera: Sapygidae). Zootaxa 3934(3):439-444. doi: https://doi.org/10.11646/zootaxa.3994.3.8 [ Links ]

Fernández F. 2000. Sistemática y filogenia de los himenópteros de la región neotropical: estado de conocimiento y perspectivas. In: Martín-Piera F, Morrone J. Melic A, editors. Hacia un proyecto CYTED para el inventario y estimación de la diversidad entomológica en Iberoamerica. Monografías Tercer Milenio, 1. Zaragoza: Sociedad Entomológica Aragonesa. p.211-231. [ Links ]

Fernández-Flores S, Fernández-Triana J, Martinez JJ, Zaldivar Riverón A. 2013. DNA barcoding species inventory of Microgastrinae wasps (Hymenoptera, Braconidae) from a Mexican tropical dry forest. Mol. Ecol. Res. 13:1146-1150. [ Links ]

Fontal-Cazalla FM, Buffington ML, Nordlander G, Liljeblad J, Ros-Farré P, Nieves-Aldrey JL, Pujade-Villar J, Ronquist F. 2002. Phylogeny of the Eucoilinae (Hymenoptera: Cynipoidea: Figitidae). Cladistics 18:154-199. doi: https://doi.org/10.1111/j.1096-0031.2002.tb00147.x [ Links ]

Forbes AA, Bagley RK, Beer MA, Hippee AC, Widmayer HA. 2018. Quantifying the unquantifiable: why Hymenoptera, not Coleoptera, is the most speciose animal order. BMC Ecol. 18:21. doi:https://doi.org/10.1186/s12898-018-0176-x [ Links ]

Gauthier N, La Salle J, Quicke DLJ, Godfray HCJ. 2000. Phylogeny of Eulophidae (Hymenoptera: Chalcidoidea), with a reclassification of Eulophinae and the recognition that Elasmidae are derived eulophids. Syst. Entomol. 25:521-539. doi: https://doi.org/10.1046/j.1365-3113.2000.00134.x [ Links ]

Genise J. 1984. Las Anthoboscinae neotropicales y comentarios sobre la diversidad de la subfamilia. Rev. Soc. Entomol. Argent. 43:195-220. [ Links ]

Genise JF. 1986. The Bradynobaenidae and some modifications to the general classification of Hymenoptera Aculeata. Physis C 44:39-54. [ Links ]

Gibson GAP, Read J, Huber JT. 2007. Diversity, classification and higher relationships of Mymarommatoidea (Hymenoptera). J. Hymenopt. Res. 16:51-146. [ Links ]

Gibson GAP. 1989. Phylogeny and classification of Eupelmidae, with a revision of the world genera of Calosotinae and Metapelmatinae (Hymenoptera: Chalcidoidea). Mem. Entomol. Soc. Can. 149:1-121. doi: https://doi.org/10.4039/entm121149fv [ Links ]

Gibson GAP. 1995. Parasitic wasps of the subfamily Eupelminae: classification and revision of World genera (Hymenoptera: Chalcidoidea: Eupelmidae). Mem. Entomol. Int. 5:1-421. [ Links ]

Gordh G, Móczár L. 1990. A catalog of the world Bethylidae (Hymenoptera, Aculeata). Mem. Am. Entomol. Inst. 46:1-364. [ Links ]

Grissell E. 1999. Hymenopteran biodiversity: some alien notions. Amer. Entomol. 45:236-244. doi: https://doi.org/10.1093/ae/45.4.235 [ Links ]

Hanson P. 1992. The Nearctic species of Ormyrus Westwood (Hymenoptera: Chalcidoidea: Ormyridae). J. Nat. Hist. 26:13331365. doi: https://doi.org/10.1080/00222939200770761 [ Links ]

Hansson C. 2009. Catalogue of the Eulophidae in the Neotropical Region. Private publication. [ Links ]

Hayat M. 1983. The genera of Aphelinidae (Hymenoptera) of the World. Syst. Entomol. 8:63-102. doi: https://doi.org/10.1111/j.1365-3113.1983.tb00467.x [ Links ]

Heraty JM. 2002. A revision of the genera of Eucharitidae (Hy-menoptera: Chalcidoidea) of the World. Mem. Amer. Entomol. Inst. Mem. Am. Entomol. Inst. 68: 1-367. [ Links ]

Huber JT. 1986. Systematics, biology, and hosts of the Mymaridae and Mymarommatidae (Insecta: Hymenoptera). Entomography 4:185-243. [ Links ]

Huber JT. 2017. Chapter 12. Biodiversity of Hymenoptera. In: Foottit RG, Adler PH, editors. Insect biodiversity: science and society. Second Edition. Estados Unidos: John Wiley & Sons Ltd. p.419-461. doi: https://doi.org/10.1002/9781118945568.ch12 [ Links ]

Johnson NF, Musetti L. 1999. Revision of the proctotrupoid genus Pelecinus Latreille (Hymenoptera: Pelecinidae). J. Nat. Hist. 33:1513-1543. doi: https://doi.org/10.1080/002229399299879 [ Links ]

Johnson NF, Musetti L. 2004. Catalog of the systematic literature of the superfamily Ceraphronoidea (Hymenoptera). Contr. Amer. Entomol. Instit. 33(2):1-149. [ Links ]

Johnson NF, Musetti L. 2012. Genera of the parasitoid wasp family Monomachidae (Hymenoptera: Diaprioidea). Zootaxa 3188(1):31-41. doi:https://doi.org/10.11646/zootaxa.3188.1.2 [ Links ]

Johnson NF. 1992. Catalog of world species of Proctotrupoidea, exclusive of Platygastridae (Hymenoptera). Mem. Am. Entomol. Inst. 51:1-835. [ Links ]

Kempf WW. 1972. Catálogo abreviado das formigas da região Neotropical. Stud. Entomol. 15:3-344. [ Links ]

Kim J-W, Heraty J. 2012. A phylogenetic analysis of the genera of Aphelininae (Hymenoptera: Aphelinidae), with generic key and descriptions of new taxa. Syst. Entomol. 37:497-549. doi: https://doi.org/10.1111/j.1365-3113.2012.00625.x [ Links ]

Kimsey LS, Bohart RM. 1991. The chrysidid wasps of the world. Oxford, New York, Toronto: Oxford University Press. [ Links ]

Kimsey LS. 1991. Relationships among the tiphiid wasp subfamilies (Hymenoptera). Syst. Entomol. 16:427-438. doi: https://doi.org/10.1111/j.1365-3113.1991.tb00677.x [ Links ]

Kimsey LS. 1992. Phylogenetic relations among the South American thynnine tiphiid wasps (Hymenoptera). Syst. Entomol. 17:133-144. doi: https://doi.org/10.1111/j.1365-3113.1992.tb00326.x [ Links ]

Klassa B, dos Santos CM. 2014. The man who loved flies: a biographical profile of Nelson Papavero. Zootaxa 3793(2):201-221. doi:https://doi.org/10.11646/zootaxa.3793.2.1 [ Links ]

Korotyaev BA, Konstantinov AS, Volkovitsh MG. 2017. Insect biodiversity in the Palearctic Region. Chapter 7. In: Foottit RG, Adler PH, editors. Insect biodiversity: Science and Society, Second Edition. Chichester, UK: John Wiley & Sons. doi: https://doi.org/10.1002/9781118945568.ch7 [ Links ]

LaSalle J. 1987. A revision of the New World Tanaostigmatidae (Hymenoptera: Chalcidoidea). Contr. Amer. Entomol. Instit. 23:1-181. [ Links ]

Li LF, Rasnitsyn AP, Shih CK, Labandeira CC, Buffington M, Li DQ, Ren D. 2018. Phylogeny of Evanioidea (Hymenoptera, Apocrita), with descriptions of new Mesozoic species from China and Myanmar. Syst. Entomol. 43:810-842. doi: https://doi.org/10.1111/syen.12315 [ Links ]

Macedo ACC. 2009. Generic classification for the Gasteruptiinae (Hymenoptera: Gasteruptiidae) based on a cladistic analysis, with the description of two new Neotropical genera and the revalidation of Plutofoenus Kieffer. Zootaxa 2075(1):1-32. doi: https://doi.org/10.11646/zootaxa.2075.1.1 [ Links ]

Macedo ACC. 2011. A revision of Gasteruption Latreille (Hymenoptera: Gasteruptiidae) in the Neotropical Region. Zootaxa 3030(1):1-62. doi: https://doi.org/10.11646/zootaxa.3030.L1 [ Links ]

Malagón-Aldana LA, Serna F, Smith DR. 2014. Siricidae (Hymenoptera) in Colombia, the first report of Urocerus gigas (Linnaeus) from northern South America. Proc. Entomol. Soc. Wash. 116(2):191-192. doi: https://doi.org/10.4289/0013-8797.116.2.191 [ Links ]

Marsh P, Wild A, Whitfield J. 2013. The Doryctinae (Braconidae) of Costa Rica: genera and species of the tribe Heterospilini. ZooKeys 347: 1-474. doi: https://doi.org/10.3897/zookeys.347.6002 [ Links ]

Masner L, García R JL. 2002. The genera of Diapriinae (Hymenoptera: Diapriidae) in the New World. Bull. Am. Mus. Nat. Hist. 268:1-138. doi:https://doi.org/10.1206/0003-0090(2002)268<0001:TGODHD>2.0.CO;2 [ Links ]

Masner L. 1976. A revision of the Ismarinae of the New World (Hymenoptera, Proctotrupoidea, Diapriidae). Can. Entomol. 108(11):1243-1266. doi: https://doi.org/10.4039/Ent1081243-11 [ Links ]

Masner L. 2006. Capítulo 88 Ceraphronidae. In: Fernández F, Sharkey MJ, editors. Introducción a los Hymenoptera de la región Neotropical. Colombia, Bogotá: Sociedad Colombiana de Entomología y Universidad Nacional de Colombia, Bogotá. p.787-788. [ Links ]

Meier R, Blaimer BB, Buenaventura E, Hartop E, von Rintelen T, Srivathsan A, Yeo D. 2021. A re-analysis of the data in Sharkey et al.'s (2021) minimalist revision reveals that BINs do not deserve names, but BOLD Systems needs a stronger commitment to open science. Cladistics 1-12. doi: https://doi.org/10.1111/cla.12489 [ Links ]

Melo GAR. c2020. Moure's Bee Catalogue. [Last accessed: 15 Dec 2020]. http://moure.cria.org.br/index [ Links ]

Michener CD. 2007. Bees of the world. Baltimore: Johns Hopkins University Press. [ Links ]

Moure JS, Urban D, Melo GAR. 2007. Catalogue of bees (Hymenoptera, Apoidea) in the Neotropical region; Sociedade Brasileira de Entomologia: Curitiba (Paraná), xiv+1058. [ Links ]

Mundy P. 2020. Insect Atlas 2020. Heinrich Böll Stiftung Pub. [ Links ]

Nemkov PJ, Lelej AS. 2013. A cladistic analysis and classification of the subfamily Bembicinae (Hymenoptera: Crabronidae), with a key to the genera. Zootaxa 3652(2)201-231. doi: https://doi.org/10.11646/zootaxa.3652.2.1 [ Links ]

Nonveiller G. 1990. Catalogue of the Mutillidae, Myrmosidae and Bradynobaenidae of the Neotropical Region including Mexico (Insecta: Hymenoptera), Series Hymenopterorum Catalogus (nova editio), Pars 18. Dr. W. Junk, The Hague. [ Links ]

Noyes JS. 1980. A review of the genera of Neotropical Encyrtidae (Hymenoptera: Chalcidoidea). Bull. Br. Mus. (Nat. Hist.) Entomol. 41:107-253. [ Links ]

Noyes JS. c2020. Universal Chalcidoidea database. World wide web electronic publication. [Last accessed: 22 Dec 2020]. http://www.nhm.ac.uk/chalcidoids [ Links ]

Zhang J, Heraty JM, Darling C, Robert RL, Baker JA, Torréns J, Rasplus JY, Lemmon A, Lemmon AM. 2021. Anchored phylog-enomics and a revised classification of the planidial larva clade of jewel wasps (Hymenoptera: Chalcidoidea) of the planidial larva clade of jewel wasps (Hymenoptera: Chalcidoidea). Syst. Entomol. 1-25. [ Links ]

Olmi M, Virla E, Fernández F . 2000. Las avispas Dryinidae de la Región Neotropical (Hymenoptera: Chrysidoidea). Biota Col. 1(2):141-163. [ Links ]

Olmi M . 1984. A revision of the Dryinidae (Hymenoptera). Mem. Am. Entomol. Inst. 37(1-2):1-1913. [ Links ]

Olmi M . 1986. New species and genera of Dryinidae (Hymenoptera, Chrysidoidea). Frustula Entomologica 7-8(20-21):63-105. [ Links ]

Olmi M . 1989. Supplement to the revision of the world Dryinidae (Hymenoptera-Chrysidoidea). Frustula Entomologica (N.S.) 12(25):109-395. [ Links ]

Olmi M . 1995. A revision of the world Embolemidae (Hymenoptera: Chrysidoidea). Frustula Entomologica (N.S.) 17(31):85-146. [ Links ]

Olmi M . 2005. A revision of the world Sclerogibbidae (Hymenoptera Chrysidoidea). Frustula Entomologica (N.S.) 26-27(39-40):46-193. [ Links ]

Osten T. 2005. Checkliste der Dolchwespen der Welt (Insecta: Hymenoptera: Scoliidae). Bericht der Naturforschenden Gesellschaft Augsburg 220: 1-62. [ Links ]

Pagliano G, Brothers DJ, Cambra R, Lelej AS, Lo Cascio P, Matteini Palmerini M, Scaramozzino PL, Romano WKA. 2018. Checklist of names in Mutillidae (Hymenoptera), with illustrations of selected species. Boll. Mus. Reg. Sci. Nat. Torino 36(1-2):5-425. [ Links ]

Pagliano G, Romano M. 2017. World list of all known species of Bradynobaenidae (Hymenoptera). Atti Mus. Civ. Stor. Nat. Morbegno 18:15-46. [ Links ]

Pate VSL. 1947. Neotropical Sapygidae, with a conspectus of the family (Hymenoptera, Aculeata). Acta Zool. Lill, 4:393-426. [ Links ]

Pezzini C, Köhler A. 2017. Annotated keys to the species of Megaspilidae (Hymenoptera: Ceraphronoidea) of the Neotropical Region. Entomo Brasilis 10(1)37-43. doi:https://doi.org/10.12741/ebrasilis.v10i1.660 [ Links ]

Pilgrim EM, Von Dohlen CD, Pitts JP. 2008. Molecular phylogenetics of Vespoidea indicate paraphyly of the super-family and novel relationships of its component families and subfamilies. Zool. Scr. 37(5):539-560. doi:https://doi.org/10.1111/j.1463-6409.2008.00340.x [ Links ]

Pinto JD. 2006. A review of the New World genera of Trichogrammatidae (Hymenoptera). J. Hymenopt. Res. 15:38-163. [ Links ]

Quicke DLJ. 2015. The braconid and ichneumonid parasitoid wasps. Biology, systematics, evolution and ecology. Wiley Blackwell. United Kingdom: John Wiley & Sons, Ltd. doi: https://doi.org/10.1002/9781118907085 [ Links ]

Ramirez W. 1970. Taxonomic and biological studies of Neotropical fig wasps (Hymenoptera: Agaonidae). University of Kansas. Sci. Bull. 49(1):1-44. doi: https://doi.org/10.5962/bhl.part.9196 [ Links ]

Richards OW. 1978. The social wasps of the Americas (excluding the Vespinae). London: British Museum (Natural History). [ Links ]

Roig-Alsina A. 1994. A new genus of Plumariidae, with notes on the relationships among the genera of the family (Hymenoptera, Chrysidoidea, Plumariidae). Mitt. Müench. Entomol. Ges. 84:91-96. [ Links ]

Ronquist F. 1995. Phylogeny and classification of the Liopteridae, an archaic group of cynipoid wasps (Hymenoptera). Entomol. Scand. Supplement. 46:1-74. [ Links ]

Ronquist F. 1999. Phylogeny, classification and evolution of the Cynipoidea. Zool. Scr. 28:139-164. doi:https://doi.org/10.1046/j.1463-6409.1999.00022.x [ Links ]

Sann M, Niehuis O, Peters RS, Mayer C, Kozlov A, Podsiadlowski L, Bank S, Meusemann K, Misof B, Bleidorn C, Ohl M. 2018. Phylogenomic analysis of Apoidea sheds new light on the sister group of bees. BMC Evol. Biol. 18:71. doi: https://doi.org/10.1186/s12862-018-1155-8 [ Links ]

Scobble MJ. 1999. Geometrid Moths of the World. A Catalogue. (Lepidoptera, Geometridae). Australia: CSIRO Publishing. [ Links ]

Sharkey MJ , Janzen DH, Hallwachs W, Chapman EG, Smith MA, Dapkey T, Brown A, Ratnasingham S, Naik S, Manjunath R, Perez K, Milton M, Hebert P, Shaw SR, Kittel RN, Solis MA, Metz MA, Goldstein PZ, Brown JW, Quicke DLJ, van Achterberg C, Brown BV, Burns JM. 2021. Minimalist revision and description of 403 new species in 11 subfamilies of Costa Rican braconid parasitoid wasps, including host records for 219 species. ZooKeys 1013: 1-665. doi: https://doi.org/10.3897/zookeys.1013.55600 [ Links ]

Shaw SR. 1990. Phylogeny and biogeography of the parasitoid wasp family Megalyridae (Hymenoptera). J. Biog. 17(6):569-581. doi:https://doi.org/10.2307/2845141 [ Links ]

Shaw SR. 2003. A new Cryptalyra species from Colombia (Hy-menoptera: Megalyridae). Zootaxa 248(1):1-4. doi:https://doi.org/10.11646/zootaxa.248.1.1 [ Links ]

Shih C, Feng H, Liu C, Zhao Y, Ren D. 2010. Morphology, phylogeny, evolution, and dispersal of pelecinid wasps (Hymenoptera: Pelecinidae) over 165 million years. Ann. Entomol. Soc. Am. 103(6):875-885. doi:https://doi.org/10.1603/AN09043 [ Links ]

Smith DR. 1988. A synopsis of the sawflies (Hymenoptera: Sym-phyta) of America south of the United States: introduction, Xy-elidae, Pamphiliidae, Cimbicidae, Diprionidae, Xiphydriidae, Siricidae, Orussidae, Cephidae. Syst. Entomol. 13:205-261. doi: https://doi.org/10.1111/j.1365-3113.1988.tb00242.x [ Links ]

Smith DR. 1990. A synopsis of the sawflies (Hymenoptera: Symphyta) of America south of the United States: Pergidae. Rev. Bras. Entomol. 34(1):7-200. [ Links ]

Smith DR. 1992. A synopsis of the sawflies (Hymenoptera: Symphyta) of America south of the United States: Argidae. Mem. Am. Entomol. Soc. 39:1-201. [ Links ]

Smith DR. 2001. World catalog of the family Aulacidae (Hymenoptera). Contr. Entomol. Int. 4(3):263-319. [ Links ]

Smith DR. 2003a. A synopsis of the sawflies (Hymenoptera: Sym-phyta) of America south of the United States: Tenthredinidae (Nematinae, Heterarthrinae, Tenthredininae). Trans. Am. Entomol. Soc. 129(1):1-45. [ Links ]

Smith DR. 2003b. A synopsis of the sawflies (Hymenoptera: Symphyta) of America south of the United States: Tenthredinidae (Allantinae). J. Hymenopt. Res. 12(1):148-192. [ Links ]

Smith DR. 2005. Aulacidae (Hymenoptera) of northern South America, emphasizing Colombia. Trans. Am. Entomol. Soc. 131(1-2):217-253. [ Links ]

Smith DR. 2006. Checklist of the Pergidae (Hymenoptera: Symphyta) of the Western Hemisphere, with a new genus and two new species. In: Blank SM, Schmidt S, Taeger A, editors. Goecke and Evers, Keltern, Germany: Recent Sawfly Research: Synthesis and Prospects. p. 613-626. [ Links ]

Taeger A , Blank SM , Liston AD. 2010. World catalog of Symphyta (Hymenoptera). Zootaxa 2580:1-1064. [ Links ]

Townes H, Townes M. 1966. A catalogue and reclassification of the Neotropic Ichneumonidae. Mem. Am. Entomol. Inst. 8:1-367. [ Links ]

Townes H, Townes M. 1981. A revision of the Serphidae (Hymenoptera) Mem. Am. Entomol. Inst. 32:1-541. [ Links ]

Townes H. 1977a. A revision of the Rhopalosomatidae. Contr. Amer. Entomol. Instit. 15(1):1-34. [ Links ]

Townes H. 1977b. A revision of the Heloridae (Hymenoptera). Contr. Amer. Entomol. Instit. 15(2):1-12. [ Links ]

Ulrich W. 1999. The number of species of Hymenoptera in Europe and assessment of the total number of Hymenoptera in the World. Pol. J. Ent. 68:151-164. [ Links ]

Vilhelmsen L, Smith DR, Malagón-Aldana LA. 2018. A review of the South American genera of Cimbicidae (Insecta, Hymenoptera). Eur. J. Taxon. 482:1-36. doi: https://doi.org/10.5852/ejt.2018.482 [ Links ]

Vilhelmsen L, Smith DR. 2002. Revision of the 'ophrynopine' genera Argentophrynopus gen. n., Guiglia Benson, Kulcania Benson, Ophrella Middlekauff, Ophrynon Middlekauff, Ophrynopus Konow, and Stirocorsia Konow (Hymenoptera: Orussidae). Insect Syst. Evol. 33(4):387-420. doi: https://doi.org/10.1163/187631202X00217 [ Links ]

Vlug HJ. 1995. Catalogue of the Platygastridae (Platygastroidea) of the world SPB Amsterdam, Netherlands: Academic Publishing. [ Links ]

Wagner DL, Grames EM, Forister ML, Berenbaum MR, Stopakd D. 2021. Insect decline in the Anthropocene: Death by a thousand cuts. Proc. Natl. Acad. Sci. U. S. A.118(2):e2023989118. doi: https://doi.org/10.1073/pnas.2023989118 [ Links ]

Weld LH. 1952. Cynipoidea (Hym.) 1905-1950, being a Supplement to the Dalla Torre and Kieffer monograph, the Cynipidae in Das Tierreich, Lieferung 24, 1910 and bringing the systematic literature of the world up to date, including keys to families and subfamilies and list of new generic, specific and variety names. Michigan: Privately Printed, Ann Arbor. p. 1-351. [ Links ]

Wharton RA, Marsh PM, Sharkey MJ , editors. 1997. Manual of the New World genera of the family Braconidae (Hymenoptera). Washington: Special Publication - International Society of Hymenopterists No. 1. [ Links ]

Yoshimoto CM. 1990. A review of the genera of New World Mymaridae (Hymenoptera; Chalcidoidea). Gainesville, FL, EE. UU: Sandhill Crane Press. [ Links ]

Yu DS, van Achterberg C, Horstmann K. 2012. World Ichneumonoidea 2011. Taxonomy, biology, morphology and distribution. Taxapad 2011. [DVD]. Canada. [ Links ]

Zaldívar-Riverón A, Martinez JJ, Ceccarelli FS, De Jesús Bonilla VS, Rodríguez-Pérez AC, Reséndiz Flores A, Smith MA. 2010. DNA barcoding a highly diverse group of parasitoid wasps (Braconidae: Doryctinae) from a Mexican nature reserve. Mitochondrial DNA 21(S1):18-23. [ Links ]

Zhang J, Heraty JM, Darling C, Kresslein RL, Baker AJ, Torréns J, Rasplus JY, Lemmon A, Lemmon EM. 2022. Anchored phylogenomics and a revised classification of the planidial larva clade of jewel wasps (Hymenoptera: Chalcidoidea). Syst. Entomol. 47(2): 329-353. doi: https://doi.org/10.1111/syen.12533 [ Links ]

Citación: Fernández F. 2022. On the diversity of Neotropical Hymenoptera. Caldasia 44(3):502-513.doi: https://doi.org/10.15446/caldasia.v44n3.94286

Received: April 06, 2021; Accepted: February 18, 2022; Published: March 28, 2022

^{CONFLICT OF INTEREST}

The author is not aware of any affiliations, memberships, funding, or financial holdings that might be perceived as affecting the objectivity of this account.

This is an open-access article distributed under the terms of the Creative Commons Attribution License