The wasp genus Sphex in Sub-Saharan Africa (Hymenoptera: Sphecidae)

  • Thorleif H. Dörfel Museum für Naturkunde, Invalidenstraße 43, 10115 Berlin, Germany
  • Michael Ohl Museum für Naturkunde, Invalidenstraße 43, 10115 Berlin, Germany https://orcid.org/0000-0002-3120-7915
Keywords: Sphex, Afrotropic, phylogeny, barcoding

Abstract

The Afrotropical species of the sphecid genus Sphex are revised. Forty-six taxa are recognized, of which fifteen are newly described: Sphex abbotti nivarius subsp. nov., S. comorensis sp. nov., S. hades sp. nov., S. nefrens sp. nov., S. occidentalis sp. nov., S. pseudopraedator sp. nov., S. pseudosatanas sp. nov., S. pulawskii sp. nov., S. rufoclypeatus sp. nov., S. satanas memnon subsp. nov., S. schmideggeri sp. nov., S. schoutedeni malawicus subsp. nov., S. socotrensis sp. nov., S. stadelmanni rufus subsp. nov. and S. victoria sp. nov. A new subgenus, Menkeella subgen. nov., is proposed for S. paulinierii Guérin Méneville, 1843.

Four former synonyms are resurrected: Sphex pruinosus var. haemorrhoidalis Magretti, 1898, for which the replacement name of S. feijeni nom. nov. is proposed, S. camerunicus Strand, 1916, S. cinerascens Dahlbom, 1843 and S. abbotti W. Fox, 1891.

Nine of the previously valid names are shown to be junior synonyms: Sphex haemorrhoidalis basuto (Arnold, 1947) and S. haemorrhoidalis kobrowi (Arnold, 1928) = S. umtalicus Strand, 1916; S. incomptus anonymus Leclercq, 1955 = S. nigrohirtus Kohl, 1895; S. neavei (Arnold, 1928) = S. abbotti W. Fox, 1891; S. observabilis (R. Turner, 1918) = S. ahasverus Kohl, 1890; S. rufiscutis (R. Turner, 1918) and S. mochii Giordani Soika, 1942 = S. jansei Cameron, 1910; S. rufiscutis laevigatus Arnold, 1951 = S. gaullei Berland, 1927; and S. stadelmanni integer (Arnold, 1928) = S. stadelmanni Kohl, 1895. Three previously synonymized species are transferred as new synonyms to different species: Sphex conradti Berland, 1927 = S. camerunicus Strand, 1916; S. kilimandjaroensis Cameron, 1908 = S. abbotti W. Fox, 1891; and S. nigripes var. pachyderma Strand, 1916 = S. umtalicus Strand, 1916. For three species, the actual type localities are on different continents than their labels indicate: Sphex ahasverus Kohl, 1890, S. castaneipes Dahlbom, 1843 and S. optimus F. Smith, 1856, with the first occuring in Africa instead of Australia and the other two presumably occuring in South America instead of Africa.

The occurrence of the argentatus group in Sub-Saharan Africa is confirmed, and seven additional species groups are established, with all but one of them based on apomorphic morphological traits. A hypothesis on the phylogenetic relationships among the subgenera and species groups of Sphex is presented, and an identification key for both sexes as well as a placoid-based identification table for males are presented. Using Bayesian inference and maximum likelihood analysis on mitochondrial and nuclear sequence data generated from thirty-one of the recognized taxa, the monophyly of the proposed species groups is corroborated.

References

Arnold G. 1928. The Sphegidae of South Africa. Part XI. Annals of the Transvaal Museum 12: 338–375.

Arnold G. 1947. New species of African Hymenoptera. No. 7. Occasional Papers of the National Museum of Southern Rhodesia 2 (13): 131–167.

Arnold G. 1951. Sphecidae and Pompilidae (Hymenoptera) collected by Mr. K.M. Guichard in West Africa and Ethiopia. Bulletin of the British Museum (Natural History), Entomology 2: 95–183. https://doi.org/10.5962/bhl.part.27752

Astrin J.J., Stüben P.E., Misof B., Wägele J.W., Gimnich F., Raupach M.J. & Ahrens D. 2012. Exploring diversity in cryptorhynchine weevils (Coleoptera) using distance-, character- and tree-based species delineation. Molecular Phylogenetics and Evolution 63 (1): 1–14. https://doi.org/10.1016/j.ympev.2011.11.018

Bamps P. 1982. Flore d’Afrique centrale (Zaire-Rwanda-Burundi): Repertoire des lieux de récolte. Jardin botanique national de Belgique, Meise, Belgium.

Benson D.A., Cavanaugh M., Clark K., Karsch-Mizrachi I., Lipman D.J., Ostell J. & Sayers E.W. 2013. GenBank. Nucleic Acids Research 41 (D1): D36–D42. https://doi.org/10.1093/nar/gks1195

Berland L. 1927. Les Sphegidae (Hyménoptères) du Muséum National de Paris (4e note). Bulletin du Muséum national d’Histoire naturelle 33: 150–156.

Berland L. 1952. La réserve naturelle integrale du Mt. Nimba. XII. – Hyménoptères vespiformes. Mémoires de l’Institut français d’Afrique noire 19: 271–276.

Berland L. 1956. Les Sphex africains (Hymenoptera). Bulletin de l’Institut français dʹAfrique noire, Série A, Sciences Naturelles 18 (4) : 1161–1181.

Bernstein H., Byerly H.C., Hopf F.A. & Michod R.E. 1985. Sex and the emergence of species. Journal of Theoretical Biology 117 (4): 665–690. https://doi.org/10.1016/S0022-5193(85)80246-0

Bezdek J., Hula V., Purchart L. & Kral K. 2012. List of local Socotran geographical names used in entomological literature. Acta Entomologica Musei Nationalis Pragae 52 (Suppl 2): 27–67.

Blaimer B.B., Lloyd M.W., Guillory W.X. & Brady S.G. 2016. Sequence capture and phylogenetic utility of genomic ultraconserved elements obtained from pinned insect specimens. PLoS One 11 (8): e0161531. https://doi.org/10.1371/journal.pone.0161531

Bohart R.M. & Menke A.S. 1963. A reclassification of the Sphecinae, with a revision of the Nearctic species of the tribes Sceliphronini and Sphecini. University of California Publications in Entomology 30: 91–182.

Bohart R.M. & Menke A.S. 1976. Sphecid Wasps of the World: a Generic Revision. University of California Press. https://doi.org/10.1525/9780520309548

Braby M.F., Eastwood R. & Murray N. 2012. The subspecies concept in butterflies: has its application in taxonomy and conservation biology outlived its usefulness? Biological Journal of the Linnean Society 106: 699–716. https://doi.org/10.1111/j.1095-8312.2012.01909.x

Brauns H. 1917. Notes and synonymy of Hymenoptera in the collection of the Transvaal Museum. Annals of the Transvaal Museum 5 (4): 238–245.

Cameron P. 1908. Hymenoptera, 7. Fossores. In: Sjöstedt Y. (ed.) Wissenschaftliche Ergebnisse der schwedischen zoologischen Expedition nach dem Kilimanjaro, dem Meru und den umgebenden Massaisteppen Deutsch-Ostafrikas 1905–1906. Volume 2 (8–14): 147–296. P. Palmquists Aktiebolag, Stockholm. [cover dated 1910, but page 294 dated Mar. 1908]

Cameron P. 1910. On the aculeate Hymenoptera collected by Mr. A.J.T. Janse, Normal College, Pretoria, in the Transvaal. Annals of the Transvaal Museum 2: 116–154.

Christ J.L. 1791. Naturgeschichte, Klassification und Nomenclatur der Insekten vom Bienen, Wespen und Ameisengeschlecht: als der fünften Klasse fünfte Ordnung des Linneischen Natursystems von den Insekten, Hymenoptera: mit häutigen Flügeln. Volume 1. Hermannischen Buchhandlung, Frankfurt am Main. https://doi.org/10.5962/bhl.title.87724

Dahlbom A.G. 1843–1845. Hymenoptera Europaea praecipue borealia; Formis typicis nonnullis Specierum Generumve exoticorum aut extraneorum propter Nexum systematicus associatis; per Familias, Genera, Species et Varietates disposita atque descripta. Tomus: Sphex in sensu Linneano. Officina Lundbergiana, Lund, Sweden. https://doi.org/10.5962/bhl.title.15890

Danforth B.N., Sauquet H. & Packer L. 1999. Phylogeny of the bee genus Halictus (Hymenoptera: Halictidae) based on parsimony and likelihood analyses of nuclear EF-1α sequence data. Molecular Phylogenetics and Evolution 13 (3): 605–618. https://doi.org/10.1006/mpev.1999.0670

Danforth B.N., Brady S.G., Sipes S.D. & Pearson A. 2004. Single-copy nuclear genes recover Cretaceous-age divergences in bees. Systematic Biology 53: 309–326. https://doi.org/10.1080/10635150490423737

De Queiroz K. 2007. Species concepts and species delimitation. Systematic Biology 56 (6): 879–886. https://doi.org/10.1080/10635150701701083

De Saussure H. 1890–1892. Histoire physique, naturelle et politique de Madagascar publiée par Alfred Grandidier. Volume 20. Histoire naturelle des Hyménoptères. Imprimerie Nationale, Paris.

Dollfuss H. 2008. The Sphecini wasps of the genera Chilosphex Bohart & Menke, Isodontia Patton, Palmodes Kohl, Prionyx Vander Linden and Sphex Linnaeus of the “Biologiezentrum Linz” collection in Linz, Austria, (Hymenoptera, Apoidea, Sphecidae). Linzer Biologische Beiträge 40: 1399–1434.

Dörfel T.H. & Ohl M. 2015. A revision of the Australian digger wasps in the genus Sphex (Hymenoptera, Sphecidae). ZooKeys 521: 1–104. https://doi.org/10.3897/zookeys.521.5995

Drury D. 1770–1782. Illustrations of Natural History. Wherein are exhibited upwards of two hundred and forty figures of exotic insects, according to their different genera; very few of which have hitherto been figured by any author, being engraved and coloured from nature, with the greatest accuracy, and under the author’s own inspection, on fifty copper-plates. With a particular description of each insect: interspersed with remarks and reflections on the nature and properties of many of them. To which is added, a translation into French. B. White, London.

Fabricius J.C. 1781. Species Insectorum exhibentes eorum Differentias specificas, Synonyma Auctorum, Loca natalia, Metamorphosin adiectis Observationibus, Descriptionibus. Volume 1. C.E. Bohnii, Hamburgi et Kilonii [= Hamburg and Kiel]. https://doi.org/10.5962/bhl.title.36509

Fabricius J.C. 1787. Mantissa Insectorum sistens eorum Species nuper detectas adiectis Characteribus genericis, Differentiis specificis, Emendationibus, Observationibus. Volume 1. C.G. Proft, Hafniae [= Copenhagen]. https://doi.org/10.5962/bhl.title.36471

Fabricius J.C. 1793. Entomologia systematica emendata et aucta. Secundum Classes, Ordines, Genera, Species adjectis Synonymis, Locis, Observationibus, Descriptionibus. 4 volumes. C.G. Proft, Hafniae [= Copenhagen]. https://doi.org/10.5962/bhl.title.122153

Faircloth B.C., Branstetter M.G., White N.D. & Brady S.G. 2014. Target enrichment of ultraconserved elements from arthropods provides a genomic perspective on relationships among Hymenoptera. Molecular Ecology Resources 15 (3): 489–501. https://doi.org/10.1111/1755-0998.12328

Fernald H.T. 1907. A collection of Sphecidae from Argentine. Bulletin of the Museum of Comparative Zoology 50: 263–272.

Field J., Ohl M. & Kennedy M. 2011. A molecular phylogeny for digger wasps in the tribe Ammophilini (Hymenoptera, Apoidea, Sphecidae). Systematic Entomology 36 (4): 732–740. https://doi.org/10.1111/j.1365-3113.2011.00591.x

Fox W.J. 1891. On three species of Hymenoptera from East Africa. Entomological News 2: 42.

Folmer O., Black M., Hoeh W., Lutz R. & Vrijenhoek R. 1994. DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3: 294–299.

Germar E.F. 1817. Reise nach Dalmatien und in das Gebiet von Ragusa. F.A. Brockhaus, Leipzig and Altenburg, Germany.

Gerstaecker A. 1871. Beitrag zur Insektenfauna von Zanzibar. Archiv für Naturgeschichte 37 (1): 345– 363.

Gess S.K. & Gess F.W. 2003. A Catalog for Flower Visiting Records for Aculeate Wasps and Bees in the Semi-Arid to Arid Areas of Southern Africa. Department of Entomology, Albany Museum, Grahamstown, South Africa.

Giordani Soika A. 1942. Sfegidi (Hym.) raccolti nellʼAfrica orientale dal Prof. Alberto Mochi e dal Dott. Marcello Mochi: 1. – Nuove specie dei generi Sphex e Cerceris. Atti della Società italiana di Scienze naturali e del Museo civico di Storia naturale di Milano 81: 196–209.

Gribodo G. 1879. Note imenotterologiche. Annali del Museo civico di Storia naturale di Genova 14: 325–347.

Gribodo G. 1894. II. Aculeati e Chrisidi. In: Emery C., Gribodo G. & Kriechbaumer G. (eds) Rassegna deglʼImenott eri raccolti nel Mozambico dal Cav. Fornasini esistenti nel Museo Zoologico della R. Università di Bologna. Memorie della R. Accademia delle Science dell’Istituto di Bologna, Series 5 4: 111–156.

Gu X., Fu Y.X. & Li W.H. 1995. Maximum likelihood estimation of the heterogeneity of substitution rate among nucleotide sites. Molecular Biology and Evolution 12 (4) : 546–557.

Guérin-Méneville F.E. 1843. Description de quelques espèces d’Hyménoptères fouisseurs de l’ancien continent, avec des observations sur plusieurs Pompiles nouveaux figurés dans la grande description d’Egypte et restés jusqu’ici inconnus. Magasin de Zoologie, dʼAnatomie comparée et de Palæontologie, 2e Série 5: 1–8.

Guiglia D. 1939. Imenotteri aculeati raccolti in Eritrea e in Etiopia dal Ten. Colon. Dr. Giorgio Invrea. Memorie della Società entomologica italiana 17: 188–202.

Hafner M.S., Sudman P.D., Villablanca F.X., Spradling T.A., Demaster J.W. & Nadler S.A. 1994. Disparate rates of molecular evolution in cospeciating hosts and parasites. Science 265: 1087–1090. https://doi.org/10.1126/science.8066445

Haig S.M., Beever E.A., Chambers S.M., Draheim H.M., Dugger B.D., Dunham S., Elliott-Smith E., Fontaine J.B., Kesler D.C., Knaus B.J., Lopes I.F., Loschl P., Mullins T.D. & Sheffield L.M. 2006. Taxonomic considerations in listing subspecies under the US Endangered Species Act. Conservation Biology 20 (6): 1584–1594. https://doi.org/10.1111/j.1523-1739.2006.00530.x

Hall T.A. 1999. BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series 41: 95–98.

Hasegawa M., Kishino H. & Yano T. 1985. Dating of human-ape splitting by a molecular clock of mitochondrial DNA. Journal of Molecular Evolution 22 (2): 160–174. https://doi.org/10.1007/BF02101694

Hensen R.V. 1991. Review of Malesian Sphecina (Hymenoptera, Sphecidae, Sphecinae). Tijdschrift voor Entomologie 134: 9–30.

Kalyaanamoorthy S., Minh B.Q., Wong T.K.F., Haeseler A. von & Jermiin L.S. 2017. ModelFinder: fast model selection for accurate phylogenetic estimates. Nature Methods 14: 587–589. https://doi.org/10.1038/nmeth.4285

Kirby W.F. 1900. Descriptions of the new species of Hymenoptera. Bulletin of the Liverpool Museums under the City Council 3: 13–24.

Kohl F.F. 1890. Die Hymenopterengruppe der Sphecinen. I. Monographie der natürlichen Gattung Sphex Linné (sens. lat.). Annalen des K. K. Naturhistorischen Hofmuseums 5 (2): 77–194. Available from https://www.biodiversitylibrary.org/page/4046912 [accessed 22 Dec. 2021].

Kohl F.F. 1895. Zur Monographie der natürlichen Gattung Sphex Linné. Annalen des K. K. natur-historischen Hofmuseums 10: 42–74. https://doi.org/10.5962/bhl.part.17608

Kohl F.F. 1898. Über neue Hymenopteren. Természetrajzi Füzetek 21: 325–367.

Kohl F.F. 1906. Zoologische Ergebnisse der Expedition der kaiserlichen Akademie der Wissenschaften nach Südarabien und Sokótra im Jahre 1898-1899. Hymenopteren. Denkschriften der kaiserlichen Akademie der Wissenschaften in Wien, Mathematisch-Naturwissenschaftliche Klasse 71: 169–301.

Kohl F.F. 1909. Sphegiden und Pompiliden von Madagaskar, den Comoren und Ostafrika. In: Voeltzkow A. (ed.) Reise in Ostafrika in den Jahren 1903–1905 mit Mitteln der Hermann und Elise geb. Heckmann Wentzel-Stiftung ausgeführt. Wissenschaftliche Ergebnisse. Band 2. Systematische Arbeiten. Issue 4: 369–378. E. Schweitzerbartsche Verlagsbuchhandlung Nägele & Dr Sproesser, Stuttgart.

Kohl F.F. 1913. Neue Pompiliden und Sphegiden vom belgischen Congogebiete. Revue zoologique africaine 3: 182–209.

Leclercq J. 1955. Hymenoptera Sphecoidea (Sphecidae I. Subfam. Sphecinae). Exploration du Parc national de l’Upemba 34:1–137.

Leclercq J. 1961. Sur les Sphex africains du groupe haemorrhoidalis Fabricius (Hymenoptera Sphecidae). Bulletin de lʼInstitut agronomique de Gembloux et des Stations de Recherche de Gembloux 29: 323–327.

Lepeletier de Saint Fargeau A.L.M. 1845. Histoire naturelle des Insectes. Hyménoptères. Volume 3. Librairie Encyclopédique de Roret, Paris.

Lindahl T. 1993. Instability and decay of the primary structure of DNA. Nature 362 (6422): 709–715. https://doi.org/10.1038/362709a0

Magretti P. 1884. Risultati di raccolte imenotterologiche nell’Africa orientale. Annali del Museo civico di Storia naturale di Genova 21: 523–636.

Magretti P. 1898. Imenotteri della seconda spedizione di Don Eugenio dei Principi Ruspoli nei paesi Galla e Somali. Annali del Museo civico di Storia naturale di Genova 39: 25–56.

Mayr E. 1942. Systematics and the Origin of Species from the Viewpoint of a Zoologist. Columbia University Press, New York.

Mayr E. 1982. Of what use are subspecies? The Auk 99 (3): 593–595.

Menke A.S. 1962. Notes on two species of sphecine wasps described by H.T. Fernald from South America (Hymenoptera: Sphecidae). The Pan-Pacific Entomologist 38: 63.

Mocsáry A. 1883. Hymenoptera nova Europaea et exotica – Európai és másföldi új hártyaröpűek. Értekezések a Természettudományok Köréből 13 (11): 1–72.

Nylander J.A.A. 2004. MrModeltest v2. Program distributed by the author. Evolutionary Biology Centre, Uppsala University, Uppsala, Sweden.

Oehlke J. & Wudowenz J. 1973. Katalog der in den Sammlungen des ehemaligen Deutschen Entomologischen Institutes aufbewahrten Typen — X. Hymenoptera. Sphecidae. Beiträge zur Entomologie 23 (5–8): 421–426.

Ohl M. 1996. Die phylogenetischen Beziehungen der Sphecinae (Hymenoptera: Apoidea: “Sphecidae”) aufgrund morphologischer Merkmale des Exoskeletts. Zooloogische Beiträge (AGE) 37: 3–40.

Pérez J. 1895. Mission scientifique de M. Ch. Alluaud aux îles Séchelles (Mars-Avril-Mai 1892). 2e Mémoire. Hyménoptères. Annales de la Société entomologique de France 64: 205–212.

Posada D. & Crandall K.A. 1998. Modeltest: testing the model of DNA substitution. Bioinformatics 14 (9): 817–818. https://doi.org/10.1093/bioinformatics/14.9.817

Pulawski W.J. 2020. Catalog of Sphecidae. California Academy of Sciences, San Francisco, CA. Available from https://www.calacademy.org/scientists/projects/catalog-of-sphecidae [accessed 27 Apr. 2020].

Rogenhofer A.F. 1889. Afrikanische Schmetterlinge des k.k. naturhistorischen Hofmuseums. I. Annalen des naturhistorischen Museums in Wien 4: 547–554.

Schmid-Egger Ch. 2014. Order Hymenoptera, families Crabronidae and Sphecidae. Further records and descriptions of new species. Arthropod Fauna of the UAE 5: 521–631.

Schmid-Egger Ch., Straka J., Ljubomirov T., Blagoev G.A., Morinière J. & Schmidt S. 2018. DNA barcodes identify 99 per cent of apoid wasp species (Hymenoptera: Ampulicidae, Crabronidae, Sphecidae) from the Western Palearctic. Molecular Ecology Resources 19 (2): 476–484. https://doi.org/10.1111/1755-0998.12963

Smith F. 1873. Descriptions of new species of fossorial Hymenoptera in the collection of the British Museum, and of a species of the rare genus Iswara, belonging to the family Dorylidae. The Annals and Magazine of Natural History, Series 4 12: 253–260 (Sep.), 291–300 (Oct.), 402–415 (Oct.). https://doi.org/10.1080/00222937308680752

Souza C.A.S., Prezoto F., Lima M.S. & Pederassi J. 2015. Sexual behavior of the digger wasp Sphex ingens Smith (Hymenoptera, Sphecidae). Revista brasileira de Entomologia 59 (2): 107–112. https://doi.org/10.1016/j.rbe.2015.03.005

Strand E. 1916. Über einige exotische Crabroniden der Gattungen Sphex und Sceliphron im Deutschen Entomologischen Museum. Archiv für Naturgeschichte 81A (11): 98–109.

Swofford D.L. 2003. PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4. Sinauer Associates, Sunderland, MA.

Taschenberg E.L. 1869. Die Sphegidae des zoologischen Museums der Universität in Halle. Zeitschrift für die gesammten Naturwissenschaften 34: 407–435.

Tavaré S. 1986. Some probabilistic and statistical problems in the analysis of DNA sequences. Lectures on Mathematics in the Life Sciences, American Mathematical Society 17: 57–86.

Trifinopoulos J., Nguyen L.-T., Haeseler A. von & Minh B.Q. 2016. W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Research 44: W232–W235. https://doi.org/10.1093/nar/gkw256

Turner R.E. 1910. Additions to our knowledge of the fossorial wasps of Australia. Proceedings of the Zoological Society of London 80 (1): 253–355. https://doi.org/10.1111/j.1096-3642.1910.tb01895.x

Turner R.E. 1912. Notes on fossorial Hymenoptera. – X. On new species from the Oriental and Ethiopian Regions. The Annals and Magazine of Natural History, Series 8 10: 361–377. https://doi.org/10.1080/00222931208693249

Turner R.E. 1918. Notes on fossorial Hymenoptera. – XXXV. On new Sphecoidea in the British Museum. The Annals and Magazine of Natural History, Series 9 1: 356–364. https://doi.org/10.1080/00222931808562327

Turner R.E. 1919. Notes on fossorial Hymenoptera. – XXXVII. On some Sphecinae in the British Museum. The Annals and Magazine of Natural History, Series 9 3: 393–397. https://doi.org/10.1080/00222931908673834

Udvardy M.D.F. 1975. A Classification of the Biogeographical Provinces of the World. Volume 8. International Union for Conservation of Nature and Natural Resources, Morges, Switzerland.

Van der Vecht J. 1961. Hymenoptera Sphecoidea Fabriciana. Zoologische Verhandelingen 48: 1–85.

Van der Vecht J. 1973. Contribution to the taxonomy of the Oriental and Australian Sphecini (Hymenoptera, Sphecidae). Proceedings, Koninklijke nederlandse Akademie van Wetenschappen, Series C 76: 341–353.

Walker F. 1871. A List of Hymenopterous Insects Collected by J.K. Lord, Esq. in Egypt, in the Neighbourhood of the Red Sea, and in Arabia. With Descriptions of the New Species. E.W. Janson, London. https://doi.org/10.5962/bhl.title.8818

Yang Z. 1994. Maximum likelihood phylogenetic estimation from DNA sequences with variable rates over sites: approximate methods. Journal of Molecular Evolution 39: 306–314. https://doi.org/10.1007/BF00160154

Published
2022-02-23
How to Cite
Dörfel, T. H., & Ohl, M. (2022). The wasp genus Sphex in Sub-Saharan Africa (Hymenoptera: Sphecidae). European Journal of Taxonomy, 796(1), 1-170. https://doi.org/10.5852/ejt.2022.796.1665
Section
Monograph