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  1. Billen J, Bauweleers E, Hashim R, Ito F
    Arthropod Struct Dev, 2013 May;42(3):173-83.
    PMID: 23333930 DOI: 10.1016/j.asd.2013.01.001
    We studied the exocrine system of both workers and ergatoid queens of Protanilla wallacei using light, scanning and transmission electron microscopy. Our survey revealed the presence of 26 glands, of which 6 had never been found before in ants. Five of these represent novel discoveries for social insects in general. The overall novel discoveries comprise an epithelial stipes gland, a pharyngeal wall gland, a central petiole gland, a lateral postpetiole gland and a foot-sole gland in the hindleg pretarsi. The intramandibular epithelial gland was already reported in some bees previously, but is now for the first time also reported in ants. The exocrine system of workers and ergatoid queens is very similar, with only the spermathecal gland showing an obvious difference. This is in line with the limited anatomical as well as behavioural difference between both castes in Protanilla compared to the situation in Leptanilla.
  2. Gotoh A, Billen J, Hashim R, Ito F
    Evol. Dev., 2016 Mar;18(2):96-104.
    PMID: 26994860 DOI: 10.1111/ede.12182
    Reproductive division of labor is one of the crucial features in social insects, however, the developmental mechanisms leading to modifications in the reproductive apparatus of workers are still not very clear. Ants show a remarkable diversity in the morphological specialization of the worker's reproductive apparatus, that allows to distinguish four types, type 1: workers that have ovaries and a functional spermatheca, and that reproduce like queens, type 2: workers have ovaries and a vestigial spermatheca, type 3: workers have ovaries but no spermatheca, and type 4: workers lost both ovaries and spermatheca. We investigated morphogenesis of the worker spermatheca in 28 ant species by histological examination. In workers of type 1, the morphogenesis of the spermatheca is very similar to that in ant queens. In type 2, the spermathecal disc also differentiates, however, the development is interrupted and remains vestigial. In types 3 and 4, the absence of the spermatheca in the adult phase is caused by a degeneration after initial formation of the spermathecal disc or by a complete lack of the spermathecal discs. The timing of these interruption and degeneration events varies among species. The species exhibiting an earlier interrupting point of spermatheca formation in workers have a larger queen-worker dimorphism, that seems to be independent from ant phylogeny.
  3. Billen J, Hashim R, Ito F
    PMID: 27130260 DOI: 10.1016/j.asd.2016.04.003
    The mandibular gland in workers of the formicine ant Myrmoteras iriodum differs from other ants both in its general morphology and ultrastructural organization. The secretory cells appear in a pseudo-epithelial arrangement that gives them a clear polarity. At their apical side, the cells are characterized by a large cup-like extension of the reservoir, from which a bulbous invagination connects to a branched end apparatus. At the basal side, the cells show a labyrinth of basal invaginations, while the lateral cell contacts show clear interdigitations. The cytoplasmic composition reveals the presence of numerous round or elongate inclusions that contain crystalline material. Microtubules are abundant, and locally fibrillar regions are found. The function of the mandibular gland in M. iriodum has not yet been documented, and should be studied using gland extracts and behavioural observations.
  4. Billen J, Stroobants Z, Wenseleers T, Hashim R, Ito F
    Arthropod Struct Dev, 2013 May;42(3):165-72.
    PMID: 23333627 DOI: 10.1016/j.asd.2012.12.001
    Histological examination of serial sections through the abdomen of workers of three species of Myopias ants revealed the presence of several exocrine glands. These include the common venom and Dufour glands as well as the pygidial gland, but also more specific sternal glands and glands associated with the sting base and the gonostyli. Two of these glands have not been reported previously among ants: one is the paired oblong plate gland, that occurs next to the oblong plate and may have a pheromonal function. The other novel gland is the paired sting shaft gland, that occurs at the dorsal side in the proximal region of the sting shaft. A remarkable characteristic of these Myopias ants is that all glands of class-3 show ducts with gradually widening internal diameter. Myopias emeryi shows a clearly more simple variety of abdominal glands than Myopias maligna and M. sp.1.
  5. Ito F, Hashim R, Huei YS, Kaufmann E, Akino T, Billen J
    Naturwissenschaften, 2004 Oct;91(10):481-4.
    PMID: 15729761
    The mechanism by which palatable species take advantage of their similarity in appearance to those that are unpalatable, in order to avoid predation, is called Batesian mimicry. Several arthropods are thought to be Batesian mimics of social insects; however, social insects that are Batesian mimics among themselves are rare. In Malaysia we found a possible Batesian mimic in an arboreal ant species, Camponotus sp., which was exclusively observed on foraging trails of the myrmicine ant Crematogaster inflata. The bright yellow and black colouring pattern, as well as the walking behaviour, were very similar in both species. We observed general interactions between the two species, and tested their palatability and the significance of the remarkably similar visual colour patterns for predator avoidance. Prey offered to C. inflata was also eaten by Camponotus workers in spite of their being attacked by C. inflata, indicating that Camponotus sp. is a commensal of C. inflata. An experiment with chicks as potential predators suggests that Camponotus sp. is palatable whereas C. inflata is unpalatable. After tasting C. inflata, the chicks no longer attacked Camponotus sp., indicating that Camponotus sp. is a Batesian mimic of Crematogaster inflata.
  6. Lee CC, Nakao H, Tseng SP, Hsu HW, Lin GL, Tay JW, et al.
    Front. Zool., 2017;14:24.
    PMID: 28503187 DOI: 10.1186/s12983-017-0210-4
    BACKGROUND: Reproductive division of labor is one of the key features of social insects. Queens are adapted for reproduction while workers are adapted for foraging and colony maintenance. In many species, however, workers retain functional ovaries and can lay unfertilized male eggs or trophic eggs. Here we report for the first time on the occurrence of physogastric workers and apparent worker reproduction in the invasive yellow crazy ant Anoplolepis gracilipes (Fr. Smith). We further examined the reproductive potential and nutritional role of physogastric workers through multidisciplinary approaches including morphological characterization, laboratory manipulation, genetic analysis and behavioral observation.

    RESULTS: Egg production with two types of eggs, namely reproductive and trophic eggs, by physogastric workers was found. The reproductive egg was confirmed to be haploid and male-destined, suggesting that the workers produced males via arrhenotokous parthenogenesis as no spermatheca was discovered. Detailed observations suggested that larvae were mainly fed with trophic eggs. Along with consumption of trophic eggs by queens and other castes as part of their diet, the vital role of physogastric workers as "trophic specialist" is confirmed.

    CONCLUSION: We propose that adaptive advantages derived from worker reproduction for A. gracilipes may include 1) trophic eggs provisioned by physogastric workers likely assist colonies of A. gracilipes in overcoming unfavorable conditions such as paucity of food during critical founding stage; 2) worker-produced males are fertile and thus might offer an inclusive fitness advantage for the doomed orphaned colony.

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