Zonitis flava, copyright Didier Descouens.

Belongs within: Meloidae.

The Nemognathini are a group of blister beetles in which the tarsal claws bear ventral teeth on the dorsal blade, and male genitalia have the parameres not separated apically (Pinto & Bologna 2002).

Blister beetles
Published 9 February 2018
Zonitis sayi, copyright Carol Davis.

This is a blister beetle of the genus Zonitis. Blister beetles, the family Meloidae, get their name from their production of cantharidin, a defensive chemical that can burn the skin of would-be predators. Zonitis is a widespread genus of blister beetles with over 100 species described from around the world. However, it should be noted that its wide distribution may relate to the genus being poorly defined and future revisions may divide its members between other genera (as I believe has already happened for the Australasian ‘Zonitis‘). As it is, Zonitis species are characterised by fully developed elytra and functional wings, and cleft tarsal claws with two rows of teeth on the upper section (Enns 1956). Adult Zonitis are flower-feeders, visiting composite plants (i.e. daisies and similar plants), and some species have the mouthparts modified into a tube for sucking nectar.

Most blister beetles exhibit what is known as hypermetamorphism or hypermetaboly, where the larvae pass through morphologically differentiated stages before reaching pupation. Zonitis species develop as parasitoids or kleptoparasites of bees. Females lay large numbers of eggs (up to and exceeding 500 in a batch) on their host plant, most commonly on the flowers though sometimes on the undersides of leaves. The eggs hatch into active, long-legged larvae that attach to bees visiting the flowers and so get carried to the bee’s nest. Once there, they moult into a less mobile stage and feed on the food stores laid aside for the bee’s larva, and potentially on the larva itself. In the North American species Zonitis atripennis flavida, the beetle larva completes its development in a single cell but European species consume the contents of two bee cells before reaching maturity. Following the initial active instar, meloid larvae pass through four feeding instars before entering a quiescent, immobile stage called the hypnotheca or prepupa. The hypnotheca moults into another feeding instar before the larva finally enters the pupal stage (Bologna et al. 2008). What the point (if anything) of the hypnotheca is, I have no idea. However, it is worth noting that hypnothecae of another meloid species, Hornia boharti, have been recorded surviving for multiple years without feeding before moulting to the next instar.

Once adults emerge from the host cell, they of course disperse to conduct their own affairs. Natural history data is patchy but indications are that many species are picky in their choice of host plant. From an economic perspective, their damaging role as a parasite of pollinating bees may be partially counterbalanced by their potential role as pollinators in their own right, but who can say which way the scales lean?

Allendesalazaria nymphoides, the hidden blister beetle
Published 11 October 2021

The blister beetles of the family Meloidae have attracted attention for a number of reasons. One is their production of caustic defensive chemicals which may be powerful enough to cause severe injury to humans or their livestock. Another is their remarkable life cycles. Many blister beetles develop as nest predators or kleptoparasites of bees. The larvae of these species are hypermetamorphic with the first instar being more mobile than later stages. These mobile larvae will find bees and latch onto them so that they can be carried to the host’s nest.

Allendesalazaria nymphoides, copyright Stanislav Krejcik.

This association reaches an extreme in Allendesalazaria nymphoides of north-west Africa. This reclusive species has, to date, been recorded from localities in Morocco, Algeria and Mauritania (Bologna & Aberlenc 2002). It is readily distinguished from other blister beetles by its much-reduced elytra which are oval and widely separated from each other. It is also distinguished by claws that lack the free lower blade found in most other meloids (Bologna & Pinto 2002). Whether they produce the noxious chemicals known from other members of their family, I haven’t found a record.

Allendesalazaria nymphoides develops in the nests of solitary burrowing bees of the genus Anthophora. Adults of A. nymphoides do not feed, and never emerge from the nest in which they matured. Instead, they lay their own eggs within that same nest. Dispersal is then left to the hatching larvae that (I presume) latch onto those emerging bees that escaped their parents’ depredations. Eventually, the new generation of bees will establish nests of their own. And when they do, the blister beetles will be ready for them.

Systematics of Nemognathini
<==Nemognathini [Apalini, Zonitini]BP02
    |--Allendesalazaria Martinez de la Escalera 1910BP02
    |    `--*A. nymphoides Martinez de la Escalera 1910BP02
    |--Onyctenus Le Peletier & Audinet de Serville 1828 [=Onychoctenus Gemminger & Harold 1870]BP02
    |    `--*O. sonnerati LePeletier & Audinet-Serville 1828BP02
    |--Sitaromorpha Dokhtouroff 1890BP02
    |    `--*S. wilkinsi Dokhtouroff 1890BP02
    |--Cochliophorus Escherich 1891BP02
    |    `--*C. reitteri Escherich 1891BP02
    |--Leptopalpus Guérin de Méneville 1829–1844 [Leptopalpina]BP02
    |    `--L. rostratus (Fabricius 1792)G-PR05 (see below for synonymy)
    |--Nyadatus Aksentjev 1981BP02
    |    `--*N. telejus Aksentjev 1981BP02
    |--Sitarobrachys Reitter 1883BP02
    |    `--‘Stenoria’ thoracica Kraatz 1863 [incl. *Sitarobrachys brevipennis Reitter 1883]BP02
    |--Gnathium Kirby 1818PB02
    |    `--G. minimumPB02
    |--Rhyphonemognatha Enns 1956PB02
    |    `--R. rufaPB02
    |--Pseudozonitis Dillon 1952PB02
    |    `--P. vaurieaeBP02
    |--Ctenopus Fischer von Waldheim 1824BP02
    |    |--*C. melanogaster Fischer von Waldheim 1824BP02
    |    `--C. sinuatipennis (Fairmaire 1892)BP02
    |--Glasunovia Semenov 1895BP02
    |    |--*G. caspica Semenov 1895BP02
    |    `--G. sillemiBP02
    |--Hornia Riley 1877 [incl. Leonia Dugés 1889 non Baird 1850, Leonidia Cockerell 1900; Horniina]PB02
    |    |--H. giganteaR35
    |    `--H. minutipennisA71
    |--Tricrania LeConte 1860 [incl. Tricranioides Wellman 1910]PB02
    |    |--T. sanguinipennisPB02
    |    `--‘Tricraniodes’ stansburyiK63
    |--Megatrachelus Motschulsky 1845BP02
    |    |--*M. politus (Gebler 1832) [=Zonitis polita; incl. M. megacephalus Kono 1936]BP02
    |    |--M. pallidipennis Motschulsky 1845BP02
    |    `--‘Zonitis’ quadricollis Fairmaire 1892BP02
    |--Sitaris Latreille 1802 [incl. Filalia Martinez de la Escalera 1906]BP02
    |    |--S. muralis (Foerster 1771)G-PR05 (see below for synonymy)
    |    |--*Filalia’ cerambycina Martinez de la Escalera 1906BP02
    |    |--S. rufipennis Küster 1849G-PR05
    |    `--S. solieri Pecchioli 1839G-PR05
    |--Zonitodema Péringuey 1909BP02
    |    |--*Z. viridipennis (Fabricius 1798) [=Zonitis viridipennis]BP02
    |    |--Z. hayekae Kaszab 1954BP02
    |    |--Z. posoka Wellman 1908BP02
    |    `--Z. rufipennis Pic 1939BP02
    |--Zonitomorpha Péringuey 1909BP02
    |    |--*Z. sellata (Fahraeus 1870) [=Zonitis sellata]BP02
    |    |--Z. cribripennis (Fairmaire 1895)BP02
    |    |--Z. davidis (Fairmaire 1886)BP02
    |    |--Z. dollei (Fairmaire 1888)BP02
    |    `--Z. transgressor (Per. 1899)BP02
    |--Zonitoschema Péringuey 1909 (see below for synonymy)BP02
    |    |--*Z. coccinea (Fabricius 1801) [=Lytta coccinea]BP02
    |    |--Z. gigantea (Fairm. 1894)BP02
    |    |--*Zonitoides’ megalops Fairmaire 1883 [=*Zonitopsis megalops]BP02
    |    |--‘Stenodera’ pallidissima Reitter 1908 [=*Stenoderistella pallidissima]BP02
    |    `--Z. suaveolaK63
    |--Apalus Fabricius 1775 (see below for synonymy)BP02
    |    |--*A. bimaculatus (Linnaeus 1761)BP02 [=Meloe bimaculatusBP02, Zonitis bimaculataL02]
    |    |--A. bipunctatus Germar 1817BP02
    |    |--A. guerini (Mulsant 1858)G-PR05 [=Criolis gueriniBP02, Hapalus gueriniG-PR05]
    |    |--A. haemapterus Fairmaire 1889F89
    |    |--‘Criolis’ hilaris Marseul 1879 [=*Coriologiton hilare]BP02
    |    |--A. necydaleus (Pallas 1773) [=Meloe necydalea, *Deratus necydaleus]BP02
    |    `--A. tanganyikanus Kaszab 1981BP02
    |--Stenoria Mulsant 1857 [incl. Gineremia Pardo Alcaide 1961, Sitaropsis Iablokoff-Khnzoryan 1958]BP02
    |    |--*S. apicalis (Latreille 1802)BP02, G-PR05 [=Sitaris apicalisG-PR05, Hapalus apicalisG-PR05]
    |    |--S. analis Schaum 1859 [=Apalus analis; incl. Hapalus analis var. acutipennis Fairmaire 1881]G-PR05
    |    |--‘Apalus (*Sitaropsis)’ erevanensis Iablokoff-Khnzoryan 1958BP02
    |    |--S. hesseiBP02
    |    |--S. laterimaculata (Reitter 1898)BP02
    |    |--S. saharica Pardo Alcaide 1961 [=*Gineremia saharica]BP02
    |    `--S. thakkola Shawaller 1996BP02
    |--Euzonitis Semenov 1893BP02
    |    |--*E. sexmaculata (Olivier 1789)BP02 (see below for synonymy)
    |    |--E. fulvipennis (Fab. 1792)B92 [=Zonitis fulvipennisB90]
    |    |--E. haroldi (Harold 1870) [=E. bipunctata var. haroldi, Zonitis quadripunctata var. haroldi]G-PR05
    |    |--E. paulinae (Mulsant & Rey 1858) [=Zonitis paolinae (l. c.)]G-PR05
    |    |--E. quadrimaculata (Pallas 1782)G-PR05 (see below for synonymy)
    |    |--E. rubida Ménétriès 1832BP02
    |    `--E. terminata (Abeille de Perrin 1880)G-PR05
    |--Nemognatha Illiger 1807 [=Nematognatha Gemminger & Harold 1870; incl. Gnathonemula Aksentjev 1981; Nemognathina]BP02
    |    |  i. s.: N. chrysomelina (Fabricius 1775) (see below for synonymy)G-PR05
    |    |         *Gnathonemula’ gracilis Aksentjev 1981 [incl. N. trascaspica Kaszab 1981]BP02
    |    |         N. luteaK63
    |    |         N. piazataPB02
    |    |         N. walkeri Beauregard 1890 [=N. bicolor Walk. 1866 (preoc.)]B90
    |    |--‘Zonitis’ piazata Fabricius 1798 [incl. Z. vittata Fabricius 1801, *Nemognatha vittata]BP02
    |    |--N. (Meganemognatha Enns 1956)BP02
    |    |    `--N. (*M.) lurida LeConte 1853BP02
    |    |--N. (Pauronemognatha Enns 1956)BP02
    |    |    `--N. (*P.) nigripennis LeConte 1853BP02
    |    `--N. (Pronemognatha Enns 1956)BP02
    |         `--N. (*P.) sparsa LeConte 1868BP02
    `--Zonitis Fabricius 1775BP02, BR17
         |  i. s.: Z. abdominalis [incl. Z. festiva]B90
         |         Z. aeneiventris Redtenb. 1868M86
         |         Z. angulataB35
         |         Z. annulata Macleay 1872M86
         |         Z. apicalis Macleay 1872M86
         |         Z. auricoma Escherich 1891G-PR05
         |         Z. bellieri Reiche 1860BP02
         |         Z. bipartita Fairm. 1879M86
         |         Z. bizonata Macleay 1872M86
         |         Z. cyanipennis Pascoe 1880M86
         |         Z. cylindracea Fairm. 1880M86
         |         Z. dichroa Germ. 1848M86
         |         Z. dunnianaPB02
         |         Z. fernancastroi Pardo Alcalde 1950G-PR05
         |         Z. flaviceps Waterh. 1875M86
         |         Z. flavicrus Fairm. 1880M86
         |         Z. fuscicornis Macleay 1872M86
         |         Z. immaculata (Olivier 1789) [incl. Z. biimpressa Chevrolat 1882, Z. mutica Scriba 1790]G-PR05
         |         Z. ingigacea Fairm. 1880M86
         |         Z. janthinipennis Fairm. 1880M86
         |         Z. limbipennis Fairm. 1880M86
         |         Z. lutea Macleay 1872M86
         |         Z. maculicollis [=Euzonitis maculicollis]BP02
         |         Z. maraniBP02
         |         Z. nana Ragusa 1882BP02
         |         Z. nigriventris Mots. 1845BP02
         |         Z. nigroaenea Fairm. 1879M86
         |         Z. nigroapicata Fairm. 1880M86
         |         Z. nigroplagiata Fairm. 1880M86
         |         Z. notaticollis (Kaszab 1951) [=Zonitis notaticollis]BP02
         |         Z. obscuripes Fairm. 1879M86
         |         Z. opacorufa Fairm. 1880M86
         |         Z. pallicolor Fairm. 1880M86
         |         Z. purpureipennis Waterh. 1875M86
         |         Z. ruficollis Frivaldszky 1877G-PR05
         |         Z. rugata Fairm. 1880M86
         |         Z. rugosipennis Fairm. 1879M86
         |         Z. sedilloti Fairm. 1880M86
         |         Z. seminigra Fairm. 1879M86
         |         Z. semirufa Fairm. 1880M86
         |         Z. splendida Fairm. 1879M86
         |         Z. tenuicornis Fairm. 1880M86
         |         Z. tricolor Le Guillou 1844M86
         |         Z. ventralis Fairm. 1880M86
         |         Z. violaceipennis Waterh. 1875M86
         |--*Z. (Zonitis) flava Fabricius 1775BP02 (see below for synonymy)
         |--Z. (Neozonitis Enns 1956)BP02
         |    `--Z. (*N.) bilineata Say 1817BP02
         `--Z. (Parazonitis Enns 1956)BP02
              `--Z. (*P.) vittigera (LeConte 1853) [=Nemognatha vittigera]BP02

Apalus Fabricius 1775 [=Hapalus Illiger 1801; incl. Coriologiton Marseul 1879, Criolis Mulsant 1858, Deratus Motschulsky 1848]BP02

Euzonitis quadrimaculata (Pallas 1782) [incl. Zonitis auricoma var. bombycina Escherich 1896, E. bipunctata ab. immaculata Escherich 1891, Z. quadripunctata var. korbi Escherich 1890, Z. bipunctata ab. korbi, Z. quadripunctata (Fabricius 1801), Z. quadripustulata (l. c.)]G-PR05

*Euzonitis sexmaculata (Olivier 1789)BP02 [=Apalus sexmaculatusBP02, Zonitis sexmaculataG-PR05; incl. E. sexmaculata ab. bipunctata Ragusa 1882G-PR05, E. sexmaculata ab. discolor Escherich 1891G-PR05, Zonitis fenestrata Pallas 1781G-PR05, E. sexmaculata ab. lunata Tauscher 1812G-PR05, E. sexmaculata ab. pedemontana Pick 1901G-PR05]

Leptopalpus rostratus (Fabricius 1792)G-PR05 [=Zonitis rostratusG-PR05, Nemognatha rostrataG-PR05; incl. *Leptopalpus chevrolatii Guérin de Méneville 1829–1844BP02]

Nemognatha chrysomelina (Fabricius 1775) [=Nemognata (l. c.) chrysomelina; incl. Nemognatha chrysomelina ab. nigripes Sufrian 1853]G-PR05

Sitaris muralis (Foerster 1771)G-PR05 [=Necydalea muralisBP02; incl. Necydalis humeralis Fabricius 1775BP02, *Si. humeralisBP02, Sytaris (l. c.) humeralisG-PR05, Si. muralis var. splendida Schaufuss 1861G-PR05]

*Zonitis (Zonitis) flava Fabricius 1775BP02 [incl. Z. nigricornis Dej. in Motschoulsky 1849 (n. n.)G-PR05, Z. praeusta var. nigripennis Fabricius 1794G-PR05, Z. flava ab. nigripennisG-PR05, Z. praeusta Fabricius 1792G-PR05, Z. flava ab. thoracica Laporte de Castelnau 1840G-PR05]

Zonitoschema Péringuey 1909 [incl. Zonitoides Fairmaire 1883 non Lehmann 1862, Stenoderistella Reitter 1911, Zonitopsis Wellman 1910]BP02

*Type species of generic name indicated


[A71] Askew, R. R. 1971. Parasitic Insects. Heinemann Educational Books: London.

[B90] Beauregard, M. 1890. Notes synonymiques suivantes relativement à quelques espèces de la tribu des vésicants. Annales de la Société Entomologique de France, 6e série 9: ccxii–ccxiii.

[B35] Boisduval, J. B. 1835. Voyage de Découvertes de l’Astrolabe. Exécuté par ordre du Roi, pendant les années 1826–1827–1828–1829, sous le commandement de M. J. Dumont d’Urville. Faune entomologique de l’océan Pacifique, avec l’illustration des insectes nouveaux recueillis pendant le voyage vol. 2. Coléoptères et autres ordres. J. Tastu: Paris.

Bologna, M. A., & H.-P. Aberlenc. 2002. Allendesalazaria, un nouveau genre de Meloidae pour la faune saharienne (Coleoptera). Bulletin de la Société Entomologique de France 107 (2): 191–192.

Bologna, M. A., M. Oliverio, M. Pitzalis & P. Mariottini. 2008. Phylogeny and evolutionary history of the blister beetles (Coleoptera, Meloidae). Molecular Phylogenetics and Evolution 48: 679–693.

[BP02] Bologna, M. A., & J. D. Pinto. 2002. The Old World genera of Meloidae (Coleoptera): a key and synopsis. Journal of Natural History 36 (17): 2013–2102.

[BR17] Bouchet, P., J.-P. Rocroi, B. Hausdorf, A. Kaim, Y. Kano, A. Nützel, P. Parkhaev, M. Schrödl & E. E. Strong. 2017. Revised classification, nomenclator and typification of gastropod and monoplacophoran families. Malacologia 61 (1–2): 1–526.

Enns, W. R. 1956. A revision of the genera Nemognatha, Zonitis, and Pseudozonitis (Coleoptera, Meloidae) in America north of Mexico, with a proposed new genus. University of Kansas Science Bulletin 37 (2): 685–909.

[F89] Fairmaire, L. 1889. Coléoptères de l’intérieur de la Chine. 5e partie. Annales de la Société Entomologique de France, 6e série 9: 5–84.

[G-PR05] García-París, M., & J. L. Ruiz. 2005. Bibliografía y registros Ibero-Baleares de Meloidae (Coleoptera) publicados hasta la aparición del “Catálogo sistemático geográfico de los coleópteros observados en la Península Ibérica, Pirineos propiamente dichos y Baleares” de J. M. de la Fuente (1933). Graellsia 61 (2): 225–255.

[K63] Kaszab, Z. 1963. Merkmale der Adaptation, Spezialisation, Konvergenz, Korrelation und Progression bei den Meloiden (Coleoptera). Acta Zoologica Academiae Scientiarum Hungaricae 9 (1–2): 135–175.

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

[M86] Masters, G. 1886. Catalogue of the described Coleoptera of Australia. Part IV. Proceedings of the Linnean Society of New South Wales, series 2, 1 (2): 259–380.

[PB02] Pinto, J. D., & M. A. Bologna. 2002. Meloidae Gyllenhal 1810. In: Arnett, R. H., Jr, M. C. Thomas, P. E. Skelley & J. H. Frank (eds) American Beetles vol. 2. Polyphaga: Scarabaeoidea through Curculionoidea pp. 522–529. CRC Press.

[R35] Rayment, T. 1935. A Cluster of Bees: Sixty essays on the life-histories of Australian bees, with specific descriptions of over 100 new species. Endeavour Press: Sydney.

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