Aposematism and mimicry in soft‐bodied beetles of the superfamily Cleroidea (Insecta)

Abstract

Bocakova, M., Bocak, L., Gimmel, M.L., Motyka, M. & Vogler, A.P. (2016). Aposematism and mimicry in soft-bodied beetles of the superfamily Cleroidea (Insecta). -Zoologica Scripta, 45, 9-21. The evolution of animal life strategies is among the main themes of current evolutionary biology. Checkered beetles, soft-winged flower beetles and their allies (superfamily Cleroidea), exhibit well-known aposematic colour patterns, particularly in the family Cleridae, which participate in mimicry complexes mostly with unpalatable beetles, ants and velvet ants representing a M€ ullerian-Batesian continuum. Many cleroids also exhibit attenuated hardening of cuticular layers resulting in a soft-bodied appearance. Here, a molecular phylogenetic analysis of the entire Cleroidea was performed using sequences of two nuclear and two mitochondrial loci of~4 kb total length. Inferred phylogenies were used to reconstruct ancestral colour patterns and involvement in mimicry complexes. The hypothesis of a soft-bodied ancestor of Cleridae and allies was tested. The phylogenetic analyses corroborated the expanded Cleroidea concept including Byturidae and Biphyllidae formerly classified as Cucujoidea. Character state optimization showed cryptic coloration was the ancestral state in Cleroidea, from which aposematic coloration originated several times in distant cleroid lineages. Within Cleridae, mimicry also arose from an ancestor that was cryptic, and multiple lineages that mimicked unpalatable beetles (Chrysomelidae, Meloidae, Lycidae) and stinging Hymenoptera evolved. Aposematic coloration was acquired in all major clerid lineages including Thanerocleridae, which are either the sister group of Chaetosomatidae or Cleridae. These findings suggest that mimetic traits in the clerid clade evolved at various times, possibly soon after the origin of soft-bodiedness. The adaptive value of aposematism in cleroids is likely to be enhanced in soft-bodied species, as this trait provides limited means of protection against predators, and therefore may promote the acquisition of aposematic and mimetic coloration in various ecological situations.Aposematism and mimicry in Cleroidea beetles M. Bocakova et al.

Comprehensive phylogeny of the Cleroidea (Coleoptera: Cucujiformia)

“…() and Bocakova et al . (, ), respectively, and one ingroup terminal ( Biphyllus lunatus ) was derived from GenBank. Of the 18 outgroup terminals, 15 were derived from GenBank, and three originated with this study (G0042, Perilopa Erichson; G0132, Paracucujus Sen Gupta & Crowson; G0324, Rhizonium Sharp).…”

“…Members of the superfamily exhibit a wide range of natural histories and behaviour, but the most common broad feeding types appear to be predatory (typified by Cleridae and most Trogossitidae: Trogossitinae), pollen‐ or nectar‐feeding (typified by the ‘melyrid lineage’), and fungus‐feeding (typified by much of the remaining Trogossitidae, and many smaller families). Many of the cleroids in the first two categories are pronouncedly diurnal as adults, some of which have developed bright aposematic coloration and are presumed to be mimics of other, less palatable insects (particularly in Cleridae and Malachiidae; Bocakova et al ., ). Other groups are much more cryptic and/or nocturnal in their habits (most Trogossitidae and smaller families), and may be found under bark, in fungal fruiting bodies or in rotting wood.…”

“…No phylogenetic analysis has focused on the trogossitid group, but members have either been included as outgroups in studies of other cleroid lineages (Bocakova et al ., , ; Gunter et al ., ) or in broad studies of the relationships of higher‐level taxa (Lawrence et al ., ; McKenna et al ., ; Robertson et al ., ; Zhang et al ., ). These preliminary studies have suggested that Trogossitidae is not monophyletic.…”

“…() (loci used: 18S , 28S , 16S , cox1 ), which focused on relationships within the so‐called ‘melyrid lineage’; and that of Bocakova et al . () (loci used: 18S , 28S , 16S , cox1 ), which focused on evolution of aposematism and soft‐bodiedness and included primarily melyrid lineage and clerid taxa. The present study includes all of the taxa included in the latter two studies plus many of those in the former, and to improve data congruence 18S data were generated for taxa of Gunter et al .…”

Multiple origins of sexual dichromatism and aposematism within large carpenter bees

“…Female Xylocopa-who are the armed sex capable of a potent, venomous sting (Schmidt 2016)are clearly the main bearer of the aposematic trait. This type of mimicry occurs frequently across many insect orders, for example in other Hymenoptera (bumble bees: Williams 2007; mutillid wasps: Wilson et al, 2012Wilson et al, , 2015, Lepidoptera (nymphalid butterflies: Owen et al 1994;Mallet and Gilbert 1995), or Coleoptera (cleroid beetles: Bocakova et al 2016). In Müllerian mimicry, related or unrelated unpalatable (or otherwise defended) species have converged on the same, often aposematic color pattern or body form (Mallet and Joron 1999).…”