A so called "rudimentary gall" induced by the gall midge Physemocecis hartigi on leaves of Tilia intermedia
The gall midge Physemocecis hartigi Liebel (Cecidomyiidae) induces a pustule gall on the leaves of the linden (Tilia intermedia D.C.). During larval development, the gall is inconspicuous on the host. Once the larva has left the gall (2 weeks after the eggs have been laid) the tissue desiccates and turns brown. We have found that the gall is initiated by the first instar larva, which attacks epidermal cells with sharp mandibles and kills a few precisely localized cells. Cells elongate perpendicularly to the attacked leaf lamina and eventually completely cover the larva. We did not observe cell division. The second instar larva enlarges the small gall cavity by inducing cell wall maceration between the vascular bundle sheath cells and the phloem. The larva feeds on cells that protrude from the open vascular bundles and on the palisade parenchyma. Contrary to earlier observations, we found these cells to be structurally modified into a nutritive tissue. The nutritive cells are hypertrophied and turgid, and have a centrally located enlarged nucleus and small vacuoles; the hydrated cytoplasm contains numerous concentric layers of endoplasmic reticulum, as well as many dictyosomes and autophagic structures. The presence of large pit fields in the cell wall suggests intense symplastic transport of solutes. The larva feeds by puncturing the turgid cells. These punctures may reach deeply into the cells but the cell wall is never ruptured; instead, the wall grows around the mandibles. Starch accumulates around the feeding area. Farther away, the modified leaf tissues contain phenolic substances. The pustule gall of P. hartigi is a highly specialized structure, supporting rapid larval development. The lack of conspicuity and the large larval cavity allow the larva to escape most of the parasitoids. Larval development goes almost without important host defense reactions and, therefore, is of low energy costs for the host. The wounding pattern of the first instar larva is highly species specific and shows a large degree of host adaptation.Key words: gall midge, gall, nutritive tissue, Physemocecis hartigi, Tilia intermedia.