"The noble false widow spider Steatoda nobilis is an emerging public health and ecological threat" by Clive Hambler (2020) is an appealing article discussing important aspects of the ecology and distribution of a medically significant spider, and the health concerns it raises.
By contrast to previous studies (Dunbar et al., 2018; Warell et al., 1991; Bauer et al., 2019; BBC 2013, 2018), this article, with its extensive media and scientific literature review, shows that S. nobilis (Thorell, 1875) is now an important health concern in Britain. Indeed, the author shows that the population of this spider has significantly increased, at least since 1990, in both southern Britain and Ireland where it has remained greatly under-recorded. In these areas, S. nobilis is now often the dominant spider on and in buildings, in places in which there is a high a risk of bites, some of which are likely to be severe, in humans, with these bites largely under-recorded. According to Clive Hambler "There is thus a possibility of bites being left without adequate rapid treatment and monitoring - with a low but non-trivial risk of necrosis or sepsis".
The author points that one of the reasons for the lack of awareness of the risk is that arachnologists typically have a conflict of interest between the conservation of the species they study and raising concerns about spiders. This may lead them to understate the risk. Clive Hambler therefore calls for a closer, appropriately weighted attention to the frequency and risk of bites, based on all the information available, rather than being "dismissive of the possibilities of bites and impacts simply because many media reports contain major errors or alarmism". He also argues that the British Arachnological Society’s guidance on "false widow spiders" "needs substantive revision, both in terms of the likelihood of bites and the severity of effects."
Indeed, the author demonstrates that many inaccuracies have been published (see Table 3 of his manuscript) and, for each, he provides a correction and/or an alternative opinion. At the end of this MS (see Table 4), he provides testable speculations and hypotheses. As he rightly points out, testing is very important to fuel the debate, because "It will be very difficult to get a balanced and proportionate debate and response for such a confused and emotive issue, especially with the many misleading popular reports." He also suggests that research will require interdisciplinary collaboration between experts in many domains, including pathologists, immunologists, clinicians, ecologists, arachnologists, psychologists, physiologists, climatologists and epidemiologists.
This preprint is clearly descriptive and speculative, but well-written, interesting and certainly useful in terms of a review of the biology, ecology, potential dangerousness and distribution of S. nobilis, particularly for future studies. There is no doubt that arachnologists, the medical community and the media will be interested in this article, which is intended to sound the alarm. Naturalists in general will also be interested in this manuscript because it is an original and successful attempt to increase knowledge about a particular taxon based on diverse information sources.
The structure of the MS is a bit odd, with a certain toing-and-froing between the ecology/biology/distribution of the spider and the risks, dangerousness and venom of bites, but this is not problematic, as shown by the reviews of the manuscript - three reviews (available below) were written, two by specialists in this noble false widow (Michel Dugon and another researcher who wished to remain anonymous).
Despite the controversy surrounding certain of the statements made in this article, I therefore strongly recommend it and look forward to seeing the identified research priorities addressed.

References

[1] Hambler, C. (2020). The “Noble false widow” spider Steatoda nobilis is an emerging public health and ecological threat. OSF Preprints, axbd4, ver. 4 peer-reviewed and recommended by PCI Zoology. doi: 10.31219/osf.io/axbd4
[2] Dunbar J.P., Afoullouss S., Sulpice R., Dugon M.M. (2018) Envenomation by the noble false widow spider Steatoda nobilis (Thorell, 1875) - five new cases of steatodism from Ireland and Great Britain. Clin Toxicol (Phila). 56(6):433-435. doi: 10.1080/15563650.2017.1393084
[3] Warrell D.A., Shaheen J., Hillyard P.D., Jones D. (1991) Neurotoxic envenoming by an immigrant spider (Steatoda nobilis) in southern England. Toxicon. 29(10):1263-5. doi: 10.1016/0041-0101(91)90198-Z
[4] Bauer, T., Feldmeier, S., Krehenwinkel, H., Wieczorrek, C., Reiser, N. and Dreitling, R. (2019) Steatoda nobilis, a false widow on the rise: a synthesis of past and current distribution trends. NeoBiota 42: 19–43. doi: 10.3897/neobiota.42.31582
[5] BBC (2013). False widow spider bites footballer Steve Harris. http://www.bbc.co.uk/news/uk-england-devon-24470023 Accessed 1 November 2018.
[6] BBC (2018). False widow spider infestation schools to remain shut. https://www.bbc.co.uk/news/uk-england-london-45761046 Accessed 19 December 2018.

Host-parasitoid interactions have been the focus of extensive ecological research for decades. One the of the major reasons is the importance host-parasitoid interactions play for the biological control of crop pests. Parasitoids are the main natural regulators for a large number of economically important pest insects, and in many cases they could be the only viable crop protection strategy. Parasitoids are also integral part of complex food webs whose structure and diversity display large spatio-temporal variations [1-3]. With the increasing globalization of human activities, the generalized spread and establishment of invasive species is a major cause of disruption in local community and food web spatio-temporal dynamics. In particular, the deliberate introduction of non-native parasitoids as part of biological control programs, aiming the suppression of established, and also highly invasive crop pests, is a common practice with potentially significant, yet poorly understood effects on local food web dynamics (e.g. [4]).
In their study, Muru et al. [5] took advantage of an existing biological control program focusing on the Asian chestnut gall wasp Dryocosmus kuriphilus, an invasive (and highly damaging) pest of chestnut trees. The species is currently a successful invader in many geographic regions, including southern France, where local parasitoid communities failed to provide an adequate control since its widespread establishment in 2010 [6]. In response, the non-native parasitoid species Torymus sinensis, which is highly-specific to the Asian chestnut gall wasp, was massively released in commercial chestnut orchards across several regions in France and the island of Corsica. The pest population outbreak was successfully contained, and thanks to the vast amount of host-parasitoid interaction data collected as part of the program, the authors were able to explore the effects of the large fluctuations in Asian chestnut gall wasp natural abundances on native parasitoid communities, immediately before, and up to five years following the introduction of its natural enemy T. sinensis.
Using co-occurrence and clustering analyses, Muru et al. [5] demonstrate that the invasion and the consecutive (efficient) control of the Asian chestnut gall wasp by the parasitoid T. sinensis have a significant impact on the structure of local parasitoid food webs. In particular, following decline in the Asian chestnut gall wasp’s populations, native parasitoids markedly switched to alternative hosts, most likely due to their respectively higher relative abundances. This pattern seemed to be driven by the degree of generalism in native parasitoid species. Indeed, when its abundances were still relatively high, the Asian chestnut gall wasp was primarily attacked by species capable of exploiting a broad range of hosts, while at low population densities only specialist parasitoids such as Mesolobus sericeus were able to persist and compete with the non-native T. sinensis.
The current study is important for two major reasons. First, it underscores the value of long-term species interaction data in order to understand the dynamic nature of food webs, namely their structural flexibility in response to changes in the environment or, as in this case, large fluctuation in abundances of a major pest species. In this context, biological control programs could be a great source of data for exploring long-term, large-scale dynamics of species interactions, and their use in ecological studies deserves to be further emphasized. Second, the study adds to the increasing empirical evidence that mobile generalist foragers can display adaptive, frequency-dependent switching behaviour ([1], [7]), which has been suggested to act as a key stabilizing mechanism in food webs by buffering fluctuating population dynamics at larger spatial scales ([8- 10]).
However, the timing of such buffering seems important, especially in systems such as commercial chestnut orchards. Despite their capacity to adaptively switch their foraging behaviour, the response of the native parasitoid communities to the new, unfamiliar resource was not fast enough in order to contain the primary outbreak under an appropriate damage threshold, thus requiring the introduction of the more specialized parasitoid T. sinensis. Nevertheless, based on current ecological theory, results presented by Muru et al. [5] suggest that the response of native parasitoid community to fluctuating host dynamics – i.e. shifts in parasitoid foraging behaviour based on their traits – could be predictable. This is encouraging considering the growing impact of biological invasions and insect pest outbreaks, but also the need to implement efficient, yet sustainable strategies for crop protection. Future studies would show at what extent observations by Muru et al. [5] are generalizable over longer time periods or other model systems. Noticeably, better understanding about population dynamics and interactions with the broader community of hosts available across habitats should allow to fine-tune predictions about parasitoids’ response to fluctuating resources.

References

[1] Eveleigh ES, McCann KS, McCarthy PC, Pollock SJ, Lucarotti CJ, Morin B, McDougall GA, Strongman DB, Huber JT, Umbanhowar J, Faria LDB (2007). Fluctuations in density of an outbreak species drive diversity cascades in food webs. Proc. Natl. Acad. Sci. USA 104, 16976-16981. doi: 10.1073/pnas.0704301104
[2] Tylianakis JM, Tscharntke T, Lewis OT (2007). Habitat modification alters the structure of tropical host–parasitoid food webs. Nature 445, 202-205. doi: 10.1038/nature05429
[3] Murakami M, Hirao T, Kasei A (2008). Effects of habitat configuration on host–parasitoid food web structure. Ecol. Res. 23, 1039-1049. doi: 10.1007/s11284-008-0478-0
[4] Geslin B, Gauzens B, Baude M, Dajoz I, Fontaine C, Henry M, Ropars L, Rollin O, Thébault E, Vereecken NJ (2016). Massively introduced managed species and their consequences for plant–pollinator interactions. Adv. Ecol. Res. 57, 147-199. doi: 10.1016/bs.aecr.2016.10.007
[5] Muru D, Borowiec N, Thaon M, Ris N, Viciriuc M I, Warot S, Vercken E (2020) The open bar is closed: restructuration of a native parasitoid community following successful control of an invasive pest. bioRxiv, 2019.12.20.884908, ver. 6 peer-reviewed and recommended by PCI Zoology. doi: 10.1101/2019.12.20.884908
[6] Borowiec N, Thaon M, Brancaccio L, Warot S, Vercken E, Fauvergue X, Ris N, Malausa J-C (2014). Classical biological control against the chestnut gall wasp 'Dryocosmus kuriphilus' (Hymenoptera, Cynipidae) in France. Plant Prot. Q. 29, 7-10.
[7] Bartley TJ, McCann KS, Bieg C, Cazelles K, Granados M, Guzzo MM, MacDougall AS, Tunney TD, McMeans BC (2019). Food web rewiring in a changing world. Nat. Ecol. Evol. 3, 345–354. doi: 10.1038/s41559-018-0772-3
[8] Kondoh M (2003). Foraging adaptation and the relationship between food-web complexity and stability. Science. 299, 1388-1391. doi: 10.1126/science.1079154
[9] McCann KS, Rooney N (2009). The more food webs change, the more they stay the same. Philos. Trans. R. Soc. Lond. B Biol. Sci. 364, 1789-801. doi: 10.1098/rstb.2008.0273
[10] Valdovinos FS, Ramos-Jiliberto R, garay-Narváez L, Urbani P, Dunne JA (2010). Consequences of adaptive behaviour for the structure and dynamics of food webs. Ecol. Lett. 13, 1546-1559. doi: 10.1111/j.1461-0248.2010.01535.x