Arachnologische Mitteilungen 55

34 G. Barrantes, L. Segura-Hernández, D. Solano-Brenes & P. Hanson cies that use more substantial amounts of silk to construct the cocoon web. This seems to be a widespread fine-tuning adjustment between the parasitoid larva and the spider host, since only small changes occur in the cocoon web of spiders in several families that build long-lived normal webs and are parasitized by wasps in different genera (Weng & Barrantes 2007, Eberhard 2010a, Korenko et al. 2018). The second-instar larva induced K. cyrtophoroides (when separated from its web) to construct a cocoon web that con- sisted of a sparse horizontal tangle.The design of this cocoon web is totally different from the design of the natural orb- web built by the spider, and cannot be recognized as part of the normal web. The second instar larva of H. heidyae seems to be capable of inducing the spider to build a new design of cocoon web in response to unpredictable situations. Simi- larly, Anelosimus under the influence of Zatypota solanoi was also capable of building a cocoon web from scratch with a completely new design (Eberhard 2010b). This suggests that the response of polysphinctine larvae to unexpected situations may be more flexible than previously thought. The general sequence of changes induced in this host spider by the parasitoid larva and the larva’s behaviour dif- fer to some extent from those described for other species in the Polysphincta genus group (Nielsen 1923, Eberhard 2000, Weng & Barrantes 2007). The second-stage larva H. heidyae only induces its host spider to construct a sparse tangle when the spider is removed from its web before the larva kills the spider.The larva then moults to its last instar, kills the spider and constructs the cocoon, and this sequence of events is qui- te stereotypical and apparently retained across this group of parasitoid wasps. If the spider is not removed from its web, the last instar larva secures the cocoon with some of its own silk threads to the centre of the web at the beginning of con- struction, but no further modifications are observed in the spider web. The sequence of events during cocoon construction of H. heidyae is, in general, similar to that of H. argyraphaga (Eber- hard 2000) and Zatypota petronae (Weng & Barrantes 2007), but differs in some respects. Attaching the cocoon by its up- per portion to the spider web differs from H. argyraphaga whose cocoon hangs freely from the centre of the cocoon web (Eberhard 2001), but is similar to Zatypota solani in that the cocoon is attached by its upper portion to the web of its host Anelosimus (Eberhard 2010b). Hymenoepimecis heidyae ’s manipulation of its spider host is finely tuned to the design and structure of K. cyrtophoroides ’s web. In this case changes in the cocoon web are barely percep- tible, in contrast to the cocoon web of other Hymenoepimesis species (and other species of the Polysphincta genus group) in- duced in host spiders with short-lived webs (Eberhard 2000, Gonzaga et al. 2010, Barrantes et al. 2017).Thus, strong, long- lived webs with some particular design (e.g. dry threads and dense tangles) requires only a few modifications to provide protection to the cocoon wasp (Sobczak et al. 2009). Acknowledgements We thank William Eberhard, Stanislaw Korenko and an anonymous reviewer for their valuable comments and suggestions on the ma- nuscript, and the Vicerrectoría de Investigación of the Universidad de Costa Rica (project 111-B6-A48) for providing financial support (GB). References Barrantes G, Sandoval L & Hanson P 2017 Cocoon web induced by Eruga telljohanni ( Ichneumonidae: Pimplinae) in Leucauge sp. (Tetragnathidae). – Arachnology 17: 245-247 – doi: 10.13156/ arac.2017.17.5.245 Eberhard WG 2000 The natural history and behavior of Hymenoe- pimecis argyraphaga (Hymenoptera: Ichneumonidae) a parasitoid of Plesiometa argyra (Araneae: Tetragnathidae). – Journal of Hy- menoptera Research 9: 220-240 Eberhard WG 2001 Under the influence: webs and building be- havior of Plesiometa argyra (Araneae, Tetragnathidae) when parasitized by Hymenoepimecis argyraphaga (Hymenoptera, Ichneumonidae). – Journal of Arachnology 29: 354-366 – doi: 10.1636/0161-8202(2001)029[0354:UTIWAB]2.0.CO;2 Eberhard WG 2010a Recovery of spiders from the effects of pa- rasitic wasps: implications for fine-tuned mechanisms of ma- nipulation. – Animal Behaviour 79: 375-383 – doi: 10.1016/j. anbehav.2009.10.033 Eberhard WG 2010b New types of behavioral manipulation of host spiders by a parasitoid wasp. – Psyche 2010(ID950614): 1-4 – doi: 10.1155/2010/950614 Eberhard WG 2013 The polysphinctine wasps Acrotaphus tibialis, Eruga ca. gutfreundi , and Hymenoepimecis tedfordi (Hymenoptera, Ichneumonidae, Pimplinae) induce their host spiders to build modified webs. – Annals of the Entomological Society of America 106: 652-660 – doi : 10.1603/AN12147 Gauld ID 2000The re-description of Pimpline genus Hymenoepimecis (Hymenoptera: Ichneumonidae) with a description of a plesi- omorphic new Costa Rican species. – Journal of Hymenoptera Research 9: 213-219 Gonzaga MO, Loffredo AP, Penteado-Dias AM & Cardoso JCF 2016 Host behavior modification of Achaearanea tingo (Araneae: Theridiidae) induced by the parasitoid wasp Zatypota alborhombarta (Hymenoptera: Ichneumonidae). – Entomological Science 19: 133-137 – doi: 10.1111/ens.12178 Gonzaga MO, Moura RR, Pêgo PT, Bang DL & Meira FA 2015 Changes to web architecture of Leucauge volupis (Araneae: Tetra­ gnathidae) induced by the parasitoid Hymenoepimecis jordanensis (Hymenoptera: Ichneumonidae). – Behaviour 152: 181-193 – doi: 10.1163/1568539X-00003238 Gonzaga MO & Sobczak JF 2007 Parasitoid-induced mortality of Araneus omnicolor (Araneae, Araneidae) by Hymenoepimecis sp. 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