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Animal constructions are the outcomes of complex evolutionary, behavioural, and ecological forces. A brief review of diverse animal builders, the materials used, and the functions they provide their builders is provided to develop approaches to studying faecal-based constructions and faecal-carrying in leaf beetles (  Coleoptera  :   Chrysomelidae  ). Field studies, rearing, dissections, photography, and films document shields constructed by larvae in two species in two tribes of the subfamily   Cassidinae  ,    Calyptocephala   attenuata    (Spaeth, 1919) (  Spilophorini  ), and    Cassida   sphaerula    Boheman, 1853 (  Cassidini  ). Natural history notes on an undetermined   Cassidini   species and    Stolas   cucullata    (Boheman, 1862) (Tribe   Mesomphaliini  ) outline the life cycle of tortoise beetles and explain terms. Commonly, the cassidine shield comprises exuviae onto which faeces are daubed, producing a pyramidal-shaped shield that can cover most of the body (up to the pronotum). In    Cal.   attenuata    the larval shield comprises only exuviae, while in    Cass.   sphaerula   , instar 1 initiates the shield by extending its telescopic anus to apply its own faeces onto its paired caudal processes; at each moult the exuvia is pushed to the caudal process base but remains attached, then more faeces are applied over it. The larva’s telescopic anus is the only tool used to build and repair the shield, not mouthparts or legs, and it also applies chemicals to the shield. Pupae in    Cal.   attenuata    retain part of the exuviae-only shield of instar VI, while pupae in    Cass.   sphaerula    retain either the entire 5 th  instar larval shield (faeces + all exuviae) or only the 5 th  larval exuvia. The caudal processes are crucial to shield construction, shield retention on the body, and as materials of the central scaffold of the structure. They also move the shield, though the muscular mechanism is not known. Altogether the faecal + exuviae shields may represent a unique morpho-behavioural synapomorphy for the crown-clade   Cassidinae   (10 tribes, ~ 2669 species) and may have been a key innovation in subsequent radiation. Defensive shields and domiciles may help explain the uneven radiation of chrysomelid subfamilial and tribal clades.

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﻿Architecture, construction, retention, and repair of faecal shields in three tribes of tortoise beetles (  Coleoptera  ,   Chrysomelidae  ,   Cassidinae  :   Cassidini  ,   Mesomphaliini  ,   Spilophorini  )

﻿Architecture, construction, retention, and repair of faecal shields in three tribes of tortoise beetles ( Coleoptera , Chrysomelidae , Cassidinae : Cassidini , Mesomphaliini , Spilophorini )

Architecture, construction, retention, and repair of faecal shields in three tribes of tortoise beetles ( Coleoptera , Chrysomelidae , Cassidinae : Cassidini , Mesomphaliini , Spilophorini )