Stizus

Stizus fasciatus, copyright Cor Zonneveld.

Belongs within: Stizini.

Stizus is a genus of medium-sized to large wasps found in Eurasia, North America and Africa that provision their nests with Orthoptera or mantids. Members of the genus are fairly generalised in appearance with subparallel inner eye margins (Bohart & Menke 1976).

The Stizus sand wasps
Published 4 July 2021

Some years ago, I presented a post on the sand wasps of the tribe Bembicini. Bembicini are just part of the broader range of sand wasps that have been variously classified as the Bembicidae, Bembicinae or Nyssoninae (Bohart & Menke 1976; Sann et al. 2018). Another diverse subgroup of the bembicids is the genus Stizus, of which more than 120 species are found in Eurasia, Africa and North America (but not in Australia or South America). Stizus species are relatively large wasps, getting up to 3.5 cm in length. Like Bembicini, they are often brightly coloured, black banded with yellow and/or red. They are otherwise fairly generalised in appearance: the labrum is exserted but is not remarkably long like that of bembicins, and the ocelli are not reduced (Bohart & Menke 1976).

Stizus pulcherrimus, copyright Phonon B.

The nesting behaviour of Stizus species was reviewed by Evans & O’Neill (2007). All known Stizus nests are constructed in soil and sand, sometimes in relatively damp locations such as salt marshes or near water bodies. Burrows of nests may be a foot or more deep and contain multiple cells; acessory burrows are common. Though females construct their burrows strictly single-handedly, they will often nest in clusters with other females. Polidori et al. (2008) found that this clustering behaviour in the European Stizus continuus was due to females being actively attracted to nests of other females, rather than just a side effect of limited nest sites. The most commonly used prey are various Orthoptera (grasshoppers or katydids); a handful of species instead prey on mantids. Prey are paralysed by repeated stinging before being flown back to the nest carried under the female. After the prey insect has been placed in a nest cell, the female lays an egg on its thorax. In most cases, cells are fully stocked with prey before laying, but females of S. continuus have been observed carrying fresh prey back to nests in which larvae have already hatched and begun eating.

Stizus perrisi female constructing nest, copyright David Genoud.

Mating between males and females generally occurs as the newly matured females emerge from the parent nest. Males often emerge before females and begin patrolling the nesting area, searching for females and chasing away other males. In some cases, they may begin actively digging for females emerging from burrows, and a newly emerged female may find herself surrounded by a pack of competing males. In their eagerness, males may become rather hasty: males of the Japanese S. pulcherrimus have been observed attempting to force themselves on females of the related genus Bembix!

Systematics of Stizus

Characters (from Bohart & Menke 1976): Eyes converging slightly below, at least narrower below than above; a low and somewhat double convexity below mid-ocellus; mandible dentate subapically; exposed part of labrum moderately convex, usually a fourth to a third as long as broad; male antenna without a projection on flagellomere IX; scutellum typically (but not always) with a medial, pubescent depression in the female; propodeum not compressed into a ridge posterolaterally; midtibia with two apical spurs; forewing with submarginal cell II open in front; hindwing media diverging far before cu-a; median cell with two distal appendices; precoxal sulcus usually present.

<==Stizus Latreille 1802-1803 [incl. Megalostizus Schulz 1906, Megastizus Patton 1879, Stizolarra Saussure 1887]BM76
    |--S. ruficornis (Forster 1771) (see below for synonymy)BM76
    |    |--S. r. ruficornisBM76
    |    |--S. r. eremicus Bytinski-Salz 1955BM76
    |    `--S. r. strigatus Mochi 1939BM76
    |--S. adelphus Kirby 1900BM76
    |--S. aegyptius Lepeletier 1845BM76
    |--S. aestivalis Mercet 1906BM76
    |    |--S. a. aestivalisBM76
    |    `--S. a. merceti Beaumont 1951BM76
    |--S. anchoratus Mantero 1917BM76
    |--S. anchorites Turner 1920BM76
    |--S. annulatus (Klug 1845) (see below for synonymy)BM76
    |--S. apicalis Guérin-Méneville 1844BM76
    |--S. arenarum Handlirsch 1892BM76
    |--S. arnoldi Mochi 1939BM76
    |--S. atrox (Smith 1856) [=Larra atrox; incl. L. flavomaculata Smith 1873, Stizus pentheres Handlirsch 1895]BM76
    |--S. aurifluus Turner 1916BM76
    |--S. basalis Guérin-Méneville 1844 [incl. Larra infuscata Klug 1845]BM76
    |--S. baumanni Handlirsch 1901BM76
    |--S. beccarii Mantero 1917BM76
    |--S. bensoni Arnold 1951BM76
    |--S. berlandi Arnold 1945BM76
    |--S. biclipeatus (Christ 1791) (see below for synonymy)BM76
    |--S. bipunctatus (Smith 1856) [=Larra bipunctata]BM76
    |--S. bizonatus Spinola 1838 [incl. Larra sphegiformis Klug 1845]BM76
    |--S. brevipennis Walsh 1869 [=*Megastizus brevipennis; incl. Larra brendeli Taschenberg 1875]BM76
    |--S. breyeri Arnold 1936BM76
    |--S. chrysorrhoeus Handlirsch 1892BM76
    |--S. coloratus Nurse 1903BM76
    |--S. combustus (Smith 1856) [=Larra combusta; incl. L. fuliginosa Klug 1845 non Dahlbom 1843]BM76
    |--S. continuus (Klug 1835) [=Larra continua]BM76
    |--S. delessertii Guérin-Méneville 1844BM76
    |--S. deserticus Giner Marí 1945BM76
    |--S. dewitzii Handlirsch 1892 [incl. S. erythraspis Cameron 1910, S. propodealis Bischoff 1915]BM76
    |--S. dispar Morawitz 1888BM76
    |--S. elegans Dahlbom 1845BM76
    |--S. ellenbergeri Arnold 1929BM76
    |--S. emir Handlirsch 1901BM76
    |--S. erransR13
    |--S. erythraeensis Mantero 1917BM76
    |--S. erythrocephalus (Taschenberg 1880) [=Larra erythrocephala]BM76
    |--S. euchromus Handlirsch 1892BM76
    |--S. eugeniae Gussakovskij 1935BM76
    |--S. excellens Lohrmann 1943 [incl. S. eximius Arnold 1936 non Morawitz 1894]BM76
    |--S. eximius Morawitz 1894BM76
    |--S. fasciatus (Fabricius 1781) [=Bembex fasciatus; incl. B. integra Fabricius 1804, Stizus terminalis Dahlbom 1845]BM76
    |--S. fedtschenkoi Radoszkowski 1877BM76
    |--S. ferrandii Magretti 1898BM76
    |--S. ferrugineus (Smith 1856) [=Larra ferruginea; incl. S. ritsemae Handlirsch 1895]BM76
    |--S. franzi Turner 1916 [incl. S. congoensis Mantero 1917, S. neavei Kohl 1913 non Turner 1912]BM76
    |--S. fuscatus Bingham 1897BM76
    |--S. fuscipennis (Smith 1856) [=Larra fuscipennis]BM76
    |--S. gobiensis Tsuneki 1970BM76
    |--S. gracilipes Handlirsch 1892BM76
    |--S. hamatus Arnold 1929BM76
    |--S. handlirschi Radoszkowski 1893BM76
    |--S. hebraeus Balthasar 1954BM76
    |--S. hispanicus Mocsáry 1883 [incl. S. villosus Costa 1887]BM76
    |--S. histrio Morawitz 1888BM76
    |--S. hugelii Handlirsch 1892BM76
    |--S. hyalipennis Handlirsch 1892BM76
    |--S. imperator Nurse 1903BM76
    |--S. imperialis Handlirsch 1892 [incl. S. tulbaghensis Arnold 1936]BM76
    |    |--S. i. imperalisBM76
    |    `--S. i. conspicuus Arnold 1952BM76
    |--S. iridis Dow 1942BM76
    |--S. jordanicus Lohrmann 1942BM76
    |--S. kaszabi Tsuneki 1971BM76
    |--S. kiseritzkii Gussakovskij 1928BM76
    |--S. koenigi Morawitz 1888 [incl. S. tages Kirby 1889]BM76
    |--S. kohlii Mocsáry 1883BM76
    |--S. lacteipennis Mocsáry 1883BM76
    |--S. lepidus (Klug 1845) [=Larra lepida]BM76
    |--S. lineatus (Fabricius 1793) [=Bembex lineatus]BM76
    |--S. lohrmanni Bohart in Bohart & Menke 1976 (see below for synonymy)BM76
    |--S. lughensis Magretti 1898BM76
    |--S. luteotaeniatus Gussakovskij 1933BM76
    |--S. lutescens Radoszkowski 1877BM76
    |--S. marnonis Handlirsch 1892BM76
    |--S. marshalli Turner 1912BM76
    |--S. marthae Handlirsch 1892 [incl. S. cheops Morice 1897, S. cheops Bingham 1897 non Morice 1897]BM76
    |--S. melanoxanthus (Smtih 1856) [=Larra melanoxantha]BM76
    |--S. melanurus Handlirsch 1892BM76
    |--S. melleus (Smith 1856) [=Larra mellea]BM76
    |--S. mocsaryi Handlirsch 1895BM76
    |--S. mongolicus Tsuneki 1971BM76
    |--S. multicolor Turner 1916BM76
    |--S. nadigi Roth 1933BM76
    |--S. neavei Turner 1912BM76
    |--S. nigriventris Arnold 1951BM76
    |--S. niloticus Handlirsch 1892BM76
    |--S. occidentalis Parker 1929BM76
    |--S. ocellatus Gistel 1857BM76
    |--S. orientalis Cameron 1890BM76
    |--S. pauli Mantero 1917BM76
    |--S. perrisii Dufour 1838BM76
    |    |--S. p. perrisiiBM76
    |    `--S. p. ibericus Beaumont 1962BM76
    |--S. pictus Dahlbom 1845BM76
    |--S. pluschtschevskii Radoszkowski 1888BM76
    |--S. praestans Morawitz 1893BM76
    |--S. pulcherrimus (Smith 1856) [=Larra pulcherrima]BM76
    |--S. pygidialis Handlirsch 1892BM76
    |--S. quartinae (Gribodo 1884) [=Sphecius quartinae]BM76
    |--S. raddei Handlirsch 1889BM76
    |--S. rapax Handlirsch 1892BM76
    |--S. repandusG20
    |--S. richardsi Arnold 1959BM76
    |--S. rubellus Turner 1912BM76
    |--S. rubroflavus Turner 1916BM76
    |--S. rufescens (Smith 1856) [=Larra rufescens]BM76
    |--S. rufipes (Fabricius 1804) [=Liris rufipes; incl. S. gigas Walsh & Riley 1868, S. grandis Lepeletier 1845]BM76
    |--S. rufiventris Radoszkowski 1877 [incl. S. sarmaticus Morawitz 1891]BM76
    |    |--S. r. rufiventrisBM76
    |    `--S. r. compar Handlirsch 1892BM76
    |--S. rufocinctus Dahlbom 1845BM76
    |--S. rufoniger Mochi 1939BM76
    |--S. saharae Roth 1934BM76
    |--S. saussurei Handlirsch 1895BM76
    |--S. savignyi Spinola 1838 [incl. Larra succinea Klug 1845]BM76
    |--S. schmiedeknechti Handlirsch 1898BM76
    |--S. scoliaeformis Arnold 1929BM76
    |--S. scolioides Kokujev 1902BM76
    |--S. sexfasciatus (Fabricius 1793) [=Bembex sexfasciatus]BM76
    |--S. spectrum Handlirsch 1901BM76
    |--S. spinulosus Radoszkowski 1876BM76
    |--S. stevensoni Arnold 1936BM76
    |--S. storeyi Turner 1920BM76
    |--S. tenuicornis (Smith 1856) [=Larra tenuicornis]BM76
    |--S. texanus Cresson 1872BM76
    |--S. transcaspicus Radoszkowski 1893BM76
    |--S. tricolor Handlirsch 1892BM76
    |--S. tridensMS01
    |--S. tridentatus [=Bembex tridentata]L02
    |--S. tunetanus Costa 1893BM76
    |--S. vespiformis (Fabricius 1775) [=Sphex vespiformis, *Stizolarra vespiformis]BM76
    |--S. vespoides (Walker 1871) [=Larra vespoides; incl. L. argentea Taschenberg 1875, L. magnifica Smith 1873]BM76
    |--S. wheeleri Arnold 1959BM76
    |--S. zhelochovtzevi Gussakovskij 1952BM76
    |--S. zimini Gussakovskij 1928BM76
    |--S. zonatus (Klug 1845) [=Larra zonata; incl. L. dimidiata Taschenberg 1875 non Panzer 1806-1809]BM76
    `--S. zonosoma Handlirsch 1895BM76

Stizus annulatus (Klug 1845) [=Larra annulata; incl. S. eversmanni Radoszkowski 1877, S. kisilkumii Radoszkowski 1877, S. picticornis Morawitz 1891, Larra subapicalis Walker 1871]BM76

Stizus biclipeatus (Christ 1791) [=Vespa biclipeata; incl. Larra cingulata Fabricius 1798, L. nubilipennis Smith 1856, Tiphia ruficornis Fabricius 1787 nec Vespa ruficornis Forster 1771 nec Bembex ruficornis Fabricius 1787]BM76

Stizus lohrmanni Bohart in Bohart & Menke 1976 [=S. arnoldi Lohrmann 1943 non Mochi 1939; incl. S. pubescens Arnold 1929 non Klug 1835]BM76

Stizus ruficornis (Forster 1771) [=Vespa ruficornis; incl. S. distinguendus Handlirsch 1901, V. gaditana Gmelin 1790, S. komarovi Radoszkowski 1888, S. ornatus Dahlbom 1845, Larra pubescens Klug 1835, Bembex ruficornis Fabricius 1787 non Vespa ruficornis Forster 1771, *Stizus ruficornis, S. scutellaris Kirby 1900]BM76

*Type species of generic name indicated

References

[BM76] Bohart, R. M., & A. S. Menke. 1976. Sphecid Wasps of the World. University of California Press: Berkeley.

Evans, H. E., & K. M. O’Neill. 2007. The Sand Wasps: Natural History and Behavior. Harvard University Press.

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[MS01] Mocsáry, A., & V. Szépligeti. 1901. Hymenopterák [Hymenopteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 121–169. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

Polidori, C., P. Mendiola, J. D. Asís, J. Tormos, J. Selfa & F. Andrietti. 2008. Female-female attraction influences nest establishment in the digger wasp Stizus continuus (Hymenoptera: Crabronidae). Animal Behaviour 75: 1651–1661.

[R13] Reuter, O. M. 1913. Lebensgewohnheiten und Instinkte der Insekten bis zum Erwachen der sozialen Instinkte. R. Friedländer & Sohn: Berlin.

Sann, M., O. Niehuis, R. S. Peters, C. Mayer, A. Kozlov, L. Podsiadlowski, S. Bank, K. Meusemann, B. Misof, C. Bleidorn & M. Ohl. 2018. Phylogenomic analysis of Apoidea sheds new light on the sister group of bees. BMC Evolutionary Biology 18: 71.

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