Review Article

Preemptive Circular Defence of Immature Insects: Definition and Occurrences of Cycloalexy Revisited

Table 1

Reported cases of cycloalexy and how they fit within the revised definition.

OrganismCriteria Aggregation is defensive (not for nest protection) Is the behaviour cycloalexy? Reference
123
Circular formationBest defended extremity at the peripheryDefault rest position

Coleoptera: Chrysomelidae: Galerucinae
Coelomera spp.; for example, C.  ruficornis Baly, 1865; C.  helenae Jolivet, 1987; C.  raquia Bechyně, 1956; and so forthYesYes, abdomensYesYesYes[1, 58]
Dircema spp.Not observedNot observedNot observedNot observedNone observed[6, 7]

Coleoptera: Chrysomelidae: Criocerinae
Lema sp.; Lema  apicalis Lacordaire, 1845 and L.  reticulosa Clark, 1866YesYes, headsYesYesYes(Figure 1(a)) [9, 10]
Lilioceris  nigropectoralis (Pic, 1928), L.  formosana Heinze, 1943YesYes, headsYesYesYes(Figure 1(b)) [11]

Coleoptera: Chrysomelidae: Chrysomelinae
Agrosteomela  chinensis (Weise, 1922)Not observedNot observedNot observedNot observedNone observed[1113]
Chrysophtharta  obovata (Chapuis, 1877) YesYes, abdomensYesProbablyYes[11, 14]
Doryphora  paykulli (Stål, 1859), D.  reticulata Fabricius, 1787YesUnclear, abdomensYesYesNo*[15]
Eugonycha  melanostoma (Stål, 1859)YesUnclear, abdomensNot reportedNot reportedTentatively[7, 11]
Gonioctena  sibirica Kimoto, 1994RoughlyUnclear, mostly abdomensMostlyUnclearUnclear[11, 12, 16]
Labidomera  suturella Guérin-Méneville, 1838Not observedNot observedNot observedNot observedNone observed[11, 1719]
Paropsis spp.; for example, P.  aegrota Boisduval, 1835, P.  maculata (Marsham, 1908), P.  atomaria Olivier, 1807 and P.  tasmanica Baly, 1864Not circularNo, mixed extremitiesNoPerhaps, unclearNo[7, 11, 14, 20, 21]
Paropsisterna spp.Not reportedNot reportedNot reportedNot reportedNot enough information[11, 12, 14, 20]
Plagiodera spp. for example, P.  versicolora (Laicharting, 1781)Not circular
No, mixed extremities
No
No
No
[7, 2225]
Phratora spp.Not observedNot observedNot observedNot observedNone observed[7, 11]
Phyllocharis  undulata (Linnaeus, 1763)RoughlyUnclear, mostly abdomensNoNot observedNo[11, 26]
Platyphora  selva Daccordi, 1993, P.  microspina (Bechyně, 1954)YesUnclear, abdomensYesYesNo*[15, 27]
Platyphora  conviva (Stål, 1858), P.  anastomozans (Perty, 1832), P.  nigronotata (Stål, 1857), P.  nitidissima (Stål, 1857) P.  fasciatomaculata (Stål, 1857), P.  vinula (Stål, 1858)YesYes, headsYesYesYes[7, 9, 10, 28]
Proseicela  vittata (Fabricius, 1781), P.  bicruciata Jacoby, 1880, P.  spectabilis (Baly, 1858)YesUnclear, abdomensYesYesNo*(Figure 5(a)) [15]
Proseicela  crucigera (Sahlberg, 1823)YesUnclear, abdomensYesYesNot enough information[7, 9]
Pterodunga  mirabile Daccordi, 2000YesUnclear, abdomensNot reportedNot reportedNot enough information[11, 12, 19]

Coleoptera: Chrysomelidae: Cassidinae
Acromis  sparsa (Boheman, 1854)YesYes, abdomensYesYesYes*[7, 11, 29]
Aspidomorpha  puncticosta Boheman, 1854, A.  miliaris (Fabricius, 1775)YesYes, abdomensYesYesYes[7, 3032]
Chelymorpha  informis Boheman, 1854, C.  alternans Boheman, 1854, C.  cribraria (Fabricius, 1875)YesYes, abdomensYesYesYes[7, 8]
Cistudinella  foveolata (Champion, 1894)YesYes, abdomensYesProbablyYes(Figure 2(a))
Conchyloctenia  punctata (Fabricius, 1787)YesYes, abdomensYesYesYes[7, 30, 33]
Coptocycla  dolosa Boheman, 1855YesYes, abdomensYesProbablyYes(Figure 4)
Eugenysa  columbiana (Boheman, 1850), E.  coscaroni Viana, 1968YesYes, abdomensYesYesYes*(Figure 2(b)) [34, 35]
Paraselenis  flava (Linnaeus, 1758)YesYes, abdomensYesYesYes*[8]
Nuzonia sp.YesYes, abdomensYesYesYes(Figure 2(c))
Ogdoecosta  biannularis (Boheman, 1854)YesYes, abdomensYesYesYes[7, 11, 36]
Omaspides  tricolorata (Boheman, 1854), O.  pallidipennis (Boheman, 1854), O.  sobrina (Boheman, 1854), O.  bistriata (Boheman, 1854) and O.  convexicollis Spaeth, 1909YesYes, abdomensYesYesYes*[7, 11, 32, 35, 3739] (D. Windsor’s observations)
Physonota  alutacea Boheman, 1854YesYes, abdomensYesProbablyYes(Figure 2(d))
Polychalma  multicava (Latreille, 1821)YesYes, abdomensYesProbablyYes(Figure 2(e))
Stolas sp., Stolas  xanthospila (Champion, 1893)YesYes, abdomensYesProbablyYes(Figure 2(f)) [7]

Coleoptera: Curculionidae: Hyperinae
Phelypera  distigma (Boheman, 1842)YesYes, headsYesYesYes[24, 40, 41]

Diptera: Ceratopogonidae: Forcipomyiinae
Forcipomyia  fuliginosa (Meigen, 1818)YesYes, abdomensYesProbablyYes[1, 11, 4244]

Hemiptera
Not specifiedNot reportedNot reportedNot reportedNot reportedNot enough information[11, 12]
Ceroplastes sp. (Coccidea), Potnia sp. (Membracidae), Nephesa  rosea (Spinola, 1839) (Flatidae), Derbe sp. (Derbidae)Roughly to not circular No, mixed extremitiesNoUnclearNo[11]
Antiteuchus  tripterus (Fabricius, 1787) (Pentatomidae)YesUnclear, abdomens NoNoNo[45]
Parastrachiajaponensis (Scott, 1880) (Parastrachiidae)YesUnclear, abdomensNoNoNo[46]

Hymenoptera: Tenthredinoidea
Bergiana sp. (Cimbicidae)YesNot reportedNot reportedNot reportedNot enough information[1]
Perga  dorsalis Leach, 1817, P.  affinis Kirby, 1882 (Pergidae) YesYes, headsYesYesYes(Figure 3(a)) [7, 11, 12, 47, 48]
Pseudoperga  guerini (Westwood, 1880) (Pergidae)YesYes, headsYesYesYes[49]
Themos  olfersii (Klug, 1834) (Argidae)YesYes, headsYesYesYes[1, 50]
Dielocerus  diasi Smith, 1975 (Argidae)Not reportedNot reportedNot reportedUnclearNot enough information[1, 50]

Hymenoptera: other superfamilies
Trigona sp. (Apidae: Meliponinae)YesYes, headsNoNoNo[1, 7, 11, 51]
Adult Hymenoptera, bees (Apidae), wasps (Vespidae), Conomyrma spp. and numerous other ants (Formicidae)Some circular, some notYes, usually headsNoNoNo[11]
Apoica sp. (Vespidae: Polistinae)YesYes, headsYesNoNo[5254]
“Parasitic Hymenoptera larvae and pupae [on] their host”YesUnclear, abdomensNoNoNo[11]

Lepidoptera: Papilionidae: Papilioninae
Papilio  laglaizei Depuiset, 1877YesUnclear, headsYesNot reportedTentatively[24, 55]

Lepidoptera: Saturniidae
Hylesia spp. (Hemileucinae)UnclearNot reportedYesProbablyNot enough information[7]
Lonomia spp. (Hemileucinae)YesProbably, headsYesProbablyProbably(Figure 3(b)) [11, 56, 57]
Arsenura spp. (Arsenurinae)Not circularNo, mixed extremitiesYesProbably notNo[1, 11].

Lepidoptera: other families
Noctuidae and SphingidaeNot circularNo, mixed extremitiesNot reportedProbably notNo[11]

Neuroptera: Ascalaphidae
Ascaloptynx  furciger (McLachlan, 1891)Yes, around twigNo, mixed extremitiesNoYesNo[1, 11, 12, 58]

Thysanoptera: Phlaeothripidae
Anactinothrips  nigricornis Hood, 1936 and A.  gustaviae Mound & Palmer, 1983YesYes, abdomensYesProbablyYes(Figure 5(b)) [59]

Non-insect arthropods
Phronima  sedentaria (Forskål, 1775) (Crustacea: Amphipoda: Hyperiidea)YesUnclear, heads?YesNoNo*[24, 60, 61]
Platydesmidae, Unidentified sp. (Myriapoda)YesUnclear, abdomensNot reportedNot reportedTentatively analogous[24, 62]

Vertebrates
Some ungulates, for example, Muskoxen Ovibos  moschatus (Zimmermann, 1780)YesYes, headsNoYesNo[1, 11, 12]
Antarctic penguinsYesUnclear, backsYesNoNo[12, 63]

These taxa are maternally defended and pose a special challenge to the definitions of cycloalexy (see Section 3.2.5).