2015 article
'Cost' of proctodeal trophallaxis in extant termite individuals has no relevance in analysing the origins of eusociality
Nalepa, C. A. (2016, February). ECOLOGICAL ENTOMOLOGY, Vol. 41, pp. 27–30.
Based on patterns of life history and behaviour in both the subsocial sister group of termites (Cryptocercus cockroaches) and the initial stages of colony foundation in one-piece (=OP, nesting in their food source) wood-feeding termites, Nalepa (1988, 1994) proposed that a key proximate step in the origin of termite eusociality was the assumption of parental brood care duties by the first nutritionally independent offspring in the family (alloparents= helpers, functional workers). This behavioural transition and the consequent transfer of costs associated with parental investment from adult parents to juvenile alloparents provides a baseline for the subsequent evolution of life history characteristics specific to extant OP termites, including a sterile soldier caste, small body size, extreme phenotypic plasticity, neotenic reproduction and interdependence of colony members. The hypothesis is also consistent with the reproductive and developmental physiology of cockroaches in a nitrogen-limited environment (Nalepa, 2011a,b, 2015). A contrary view, that alloparental brood care is not a necessary prerequisite for termite eusociality is advanced by Korb et al. (2012) and Korb (2007, 2015), who observed little to no brood care behaviour in alloparents originating from established colonies of drywood termites (Cryptotermes). It is further argued that one component of brood care, proctodeal trophallaxis, would be of little or no cost to individual caregivers because the behaviour is typically a reciprocal interaction (Korb, 2007, 2008a,b, 2010a). Here I suggest that there is little evidence to support claims of the absence of brood care and its correlate, independent offspring, in extant OP termites, and the notion that proctodeal feeding is not costly to individual alloparents is based on misconceptions regarding what may be considered ‘ancestral’ in a termite.