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Independent introductions of hedgehogs to the North and South Island of New Zealand

we also report on several relocations from established populations in Canterbury (South Island) to regions on the North Island, and none in the opposite direction. We illustrate the importance of linking observational and molecular data with historical records when interpreting the introduction pathways of introduced species.


Introduction
New Zealand is the only country in the world where European hedgehogs (Erinaceus europaeus L.) have successfully established in the wild outside their native range (Brockie 1975;Long 2003). Hedgehogs were shipped to New Zealand by acclimatisation societies three times between 1869 and 1885, but the number of survivors was low (only six in total), and the only documented releases during that time were two males from the last shipment (Thomson 1922;Brockie 1975).
in the wild are reported from the 1890s (Thomson 1922;Brockie 1975;King 2005). Hedgehogs were brought over to New Zealand as pets but also disseminated throughout the country. This was done at least partly in the hope that they might control insect and mollusc pests (Brockie 1975;Long 2003). While they failed in this task, their voracious appetite for endemic skinks and eggs of ground-nesting birds has made The general consensus from historical records is that 19th century, and were absent on the North Island before the 20th century when they were thought to have been transported there from the South Island (Thomson 1922;Wodzicki 1950;Brockie 1975;Long 2003;King 2005). A recent study based et al. 2013) has questioned this view. Approximate Bayesian computation models supported only scenarios with colonisation of the North Island directly from the source population in Great (2013) suggested that ancient southern populations might have been replaced by the invasion waves from the North Island. Another possibility is that the historical record, as collected by Brockie (1975), is rather incomplete and that hedgehogs appeared on the North Island much earlier than previously thought.
Here we re-examine the introduction history of the European hedgehog to New Zealand by using primary sources of information, original documents of acclimatisation societies and, mainly, newspaper articles, which proved to be a reliable (https://paperspast.natlib.govt.nz), also a very powerful resource (Star 2014;Pipek et al. 2015aPipek et al. ,b, 2019King 2017a,b).

Methods
We collected information from published annual reports and unpublished minute books of acclimatisation societies in New Zealand and newspaper articles stored in the Papers Past archive (paperspast.natlib.govt.nz) published from 1850 to 1940. In the newspaper archive we used the search query: hedge*hog + hedge*hogs, which enabled any combination of the following strings, "hedge hog", "hedge-hog" or "hedgehog" in singular as well as plural form. We also explored less commonly used terms, hedge*pig and urchin, but this did not lead to the discovery of new articles.
We double checked this information with data included in George M. Thomson's monograph on plant and animal introductions (Thomson 1922) and other resources addressing the naturalisation of hedgehogs and other animals in New Zealand (Wodzicki 1950;Brockie 1975;Long 2003;King 2005). As George Thomson lived in Otago and took care of the survivors of the only shipment of hedgehogs of the Otago Acclimatisation Society in 1885, we consider his data (about presence, not absence) from this region, and also from Canterbury, as trustworthy.
The geographic distribution of the earliest records of hedgehogs, combined with knowledge of individual shipments, strongly Canterbury ( Fig. 1; See Appendix S1 in Supplementary Materials). We tracked down 10 shipments of hedgehogs, six to the South Island (1869-1894) and four to the North Island (1883-1896; Table 1). The mortality during shipping (when reported) was high (ranging from 50 to 97%, median 91%). Some hedgehogs must have been shipped without being noticed (or documented) by newspapers, and then released, either deliberately or accidentally, as in case of pets (Drummond 1911). Several releases, not reported by contemporary newspapers, were noted in hindsight (Table 1; Appendix S1).

South Island
Otago were apparently limited and contributed only marginally, if at all, to the successful establishment of hedgehogs. In all documented cases, only a few individuals were imported ( female and two males in its gardens in 1885, however the female died soon after and one of the males disappeared. In

North Island
Hedgehogs on the North Island were observed before any of the tracked overseas shipments transporting hedgehogs reached the region: in the 1880s, escaped pets were reported from Auckland (Auckland Star 1880) and Wellington (Evening Post 1885).
The local acclimatisation societies were less active than their southern counterparts, none of the tracked shipments though the acclimatisation society of Wellington wanted to establishment in 1871 (Lyttelton Times 1872).
The sole surviving hedgehog (of eight) shipped to Whanganui in 1883 (Manawatu Standard 1883), was followed 11 years later by a few individuals shipped to Wairarapa (Evening Post 1894). Two years later an unknown number were transported to Hawke's Bay (Daily Telegraph 1896) and (Marlborough Express 1898).
Apart from the direct shipments from Europe described above, some hedgehogs were imported from the South Island.
In 1906 (presumably -the date relates to the Christchurch exhibition), several hedgehogs were transported from Canterbury to the Manawatu-Whanganui region (Manawatu Standard 1911a;Woodville Examiner 1911;New Zealand Times 1911). Also in 1911, eight hedgehogs were transported from Canterbury to Poverty Bay (Poverty Bay Herald 1911). Finally, there was also some movement of hedgehogs within the North Island: a pair or two were transported from Wellington to Whanganui (Opunake Times 1927) and a pregnant female from Whanganui to the Taranaki region (Stratford Evening Post 1926). Discussion the knowledge of the history of hedgehog introductions in New Zealand. While some overseas shipments to the South Island were recorded by Brockie (1975), we managed to track yet unknown overseas shipments to the North Island. The list is nevertheless still not exhaustive; some hedgehogs appeared as pets in areas where no previous shipment was noted, i.e. in 1880s in Auckland and in Wellington. Overall, newspapers were especially valuable in capturing the early occurrences of hedgehogs on the North Island, which were missed by Brockie (1975). In later years, the data in Brockie (1975) and in newspapers correspond well. It is understandable that Brockie (1975) missed information on early occurrences because a substantial part of his data comes from questionnaires undertaken by Wodzicki (1950) and people remembering the early spread might have already passed away by that time. He also did not have the advantage we have in digitised newspaper articles. early shipments are untraceable, hedgehogs were introduced independently to the North Island of New Zealand well before the 20th century with at least four shipments from Europe. Some hedgehogs were also relocated in the 20th century from  (Fig. 1, Table 2). Support for this high rate of translocation of hedgehogs in New Zealand stems from genetic data that show no evidence of populations being isolated by distance By contrast, our data do not support the most likely establishment initially occurred in the North Island from where hedgehogs independently colonised other regions in the North as well as the South Island. According to newspaper articles, hedgehogs were already abundant in the South Island, especially around the big cities (Christchurch, Timaru, Dunedin) by the time they had just begun to be recorded in the North Island (Fig. 1, Appendix S1).
Our study demonstrates that thorough historical analysis can still add new insights into a relatively well studied species studies aiming at clarifying the history of animal introductions improved historical record, that can help to select the most appropriate set of introduction scenarios for rigorous testing (e.g. ABC analysis), and develop a balanced sampling design, based on the documented species spread, by collecting the material from likely sources of early introductions as well as from more distant sites.