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Rabbits have had a disastrous impact on Australian agriculture and native plants.Credit: Bettman/Getty

A genomic analysis has helped to show that Australia’s invasive rabbit population probably originated from a shipment of two dozen wild English rabbits that arrived near Melbourne on Christmas Day, 1859. The study also finds that the herd’s wild ancestry probably gave it an advantage over previous arrivals.Rabbits have invaded most of the Australian continent and have had a disastrous impact on ecosystems, threatening some 300 species of plants and animals, and causing hundreds of millions of dollars’ worth of damage to the agriculture industry a year. “That single event triggered this enormous catastrophe, ecologically and economically, in Australia,” says Francis Jiggins, an evolutionary geneticist at the University of Cambridge, UK, and study co-author.Breeding like rabbitsHistorical records suggest that the first European rabbits (Oryctolagus cuniculus) in Australia arrived in Sydney in 1788, with the first colonizers. Ships bringing rabbits continued to dock along the coast for decades, but it wasn’t until the second half of the nineteenth century that the population expanded significantly, spreading across the country at a rate of 100 kilometres a year.Historical records also suggest that the rabbit expansion followed a shipment of animals that arrived for a certain Thomas Austin at Barwon Park, southwest of what is now Melbourne. His brother had trapped them around their family property in Baltonsborough in southwest England.Joel Alves, an evolutionary geneticist at the University of Oxford, UK, and his colleagues wanted to find out whether genomic data corroborated the records. They analysed genetic data from 179 wild rabbits caught across Australia and in New Zealand, France and the United Kingdom, as well as 8 domestic rabbits of different breeds.They found that most rabbits in mainland Australia were genetically similar, with mixed wild and domestic ancestry. Australian rabbits also shared more rare alleles with rabbits from southwest England than with those from elsewhere in the United Kingdom, suggesting they originated in Baltonsborough. Looking specifically at mitochondrial DNA, which is inherited from the mother, the researchers concluded that most mainland Australian rabbits descended from about five females, introduced from Europe.The researchers also found that the rabbits’ genetic diversity declined the further from Barwon Park the animals were caught, and that alleles that are rare or absent in wild rabbits increased. The researchers say these patterns are consistent with the idea that most rabbits across Australia originated from Barwon Park. The team report their findings in the Proceedings of the National Academy of Sciences on 22 August1.“This is a very exciting paper on a very important and well-studied topic,” says Martin Nuñez, who researches ecological invasions at the University of Houston in Texas. Using genetics to understand how unwanted animal invasions start can help to predict future invasions, he says.Perfect stormOverall, the team says that the rabbits’ wild ancestry was an important factor in triggering their invasion of the continent. “Wild rabbits are different,” says Alves. They exhibit traits such as fleeing stressful environments and burrow-digging, meaning that they were probably better at evading predators and surviving in difficult terrain than are domestic rabbits, he says. Historical records suggest that Austin requested wild rabbits, and that previous arrivals were largely domestic breeds.The expansion of Australian pastoral lands and widespread suppression of predators around that time would have also helped their expansion. “It was like a perfect storm,” says Alves. “You have the right rabbits in the right place at the right time, with the right changes in the environment.”“The genetic analyses appear very sound,” says rabbit geneticist Amy Iannella, a consultant based in Adelaide, Australia. She adds that although the country’s rabbit populations probably originated in Barwon Park, their rapid expansion might have been aided by people transporting the animals to other parts of the country, where they also began spreading. Rabbits are typically communal animals that rely on shelter for survival and juveniles rarely travel further than 1 kilometre, she says. “The idea of rabbits moving fast enough at the invasion front to colonize Australia so quickly from a single release, well that feels extreme to me, given what we know about rabbit ecology.” More

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Discovery and historiography of USNM PAL V 11367With basic image enhancement tools (e.g., Adobe Photoshop), we were able to better resolve the original but faded specimen label in the collections associated with USNM PAL V 11367 (Fig. 1 and Related file 1). Specifically, we were able to make the now-faded handwritten notes legible (Fig. 1A,B), revealing critical information about the specimen. The widespread availability of image enhancement for faded fieldnotes and labels provides a new source of information for uncovering legacy issues in museum collections (e.g.21,22,23), especially in cases where locality data or collecting information cannot be well resolved.Accession files with this specimen (Related file 1) show that it was gifted from Arthur M. Ames to the United States National Museum (now the National Museum of Natural History, Smithsonian Institution) on 15 October 1925, and approved by George P. Merrill, head curator of geology from 1917 to 1929. Prior to its accession to the museum, an anonymous individual identified the tooth as belonging to Desmostylus hesperus. Forty years later, on 17 November 1965, Charles A. Repenning reidentified this specimen as Paleoparadoxia sp. (Fig. 1A,B), an assertion that was incorporated into its catalog information. According to the label, USNM PAL V 11367 was collected in the city of Corona, Riverside County, California, yet no precise information of its geological provenance was recorded. On the backside of the label, there are notes (Fig. 1B) referring to the US Geologic Survey Corona South 7.5′ quadrangle map for Riverside and Orange counties, California24. However, no geographic location, exact horizon, nor lithology was stated, and the specimen’s collector, A. M. Ames, lived in Santa Barbara, California but died on 25 August 193921,22,23.In nearly a century after its discovery, the only mention of USNM PAL V 11367 was by Panofsky25, who listed it in a catalog of desmostylian tooth specimens used as a comparative basis for a mandible restoration of the “Stanford specimen” N. repenningi. Panofsky25 identified USNM PAL V 11367 as a left m2 with six main cusps, with no additional cusps (Table 1 in25), while also stating that this specimen has “an open lake in the center of each of the seven cusps” (25: p. 103). The inconsistency of this description differs from our own, which we attribute to differences in morphological criteria or a typographic error.Geological horizon and age of USNM PAL V 11367In this paper, we refer to the “Topanga” Formation following recent studies20,26,27 of this geologic unit. This formation was originally based on a sequence of marine sandstones exposed in an anticline just west of Old Topanga Canyon in the central Santa Monica Mountains of Los Angeles County, California28. After its initial description, the name of the formation was applied to a much thicker and heterogeneous sequence of sedimentary and volcanic rocks29. Campbell et al.30 compiled the history and chronology of changes in usage of “Topanga” in the Miocene stratigraphic nomenclature in Southern California, showing that the criteria of continuous deposition and shared provenance were not demonstrated in every instance. Campbell et al.30 argued that strata assigned to the Topanga Formation in the Los Angeles Basin and eastern Ventura Basin areas are different from other units that have also been referred to the Topanga Formation in Orange County or in the Santa Monica Mountains of Los Angeles and Ventura counties. To distinguish these units, here we follow recent studies20,26,27 and use the name of “Topanga” Formation for the early to middle Miocene rocks bearing fossil marine mammals20,26,31,32,33 in Southern California.According to the collections records (Fig. 1), USNM PAL V 11367 was collected in the city of Corona, Riverside County, California, USA. This city is in the western part of Riverside County, comprising an approximate area of 100 km234. Previously, Panofsky25 suggested that USNM PAL V 11367 would have derived from the Temblor Formation (14.8 to 15.8 Ma35), likely as a guess based on the prevalence of desmostylian teeth recovered from this unit in central California, yet today there are no Temblor Formation outcrops mapped near Corona24,36; the closest Temblor outcrops are located in Fresno and Kern counties37, approximately 200 km away.The geologic maps of Riverside County24,36,38 indicate that the city limits of Corona encompass a wide variety of sedimentary rocks from the Jurassic to the Holocene in age, but only a few marine deposits, such as the Jurassic Bedford Canyon Formation and the middle Miocene “Topanga” Formation are exposed24,39. Specifically, the marine sandstones of the “Topanga” Formation occur within the fault zone at the southeast and northwest of Corona.Outside of Riverside County, the “Topanga” Formation has yielded a diverse assemblage of fossil marine vertebrates in Southern California20,26,31, including desmostylians referred to Desmostylus hesperus and Paleoparadoxia sp. in Orange County (Supplementary 1). USNM PAL V 11367 represents the second reported fossil marine mammal from Riverside County. Previously, an isolated record of “Cetacea indet.” was mentioned from the Zanclean stage Imperial Formation40 and Supplementary Data 2), which is exposed far east of Corona’s city limits.In assessing the age of the “Topanga” Formation in Southern California, Boessenecker and Churchill26,31 argued that the land mammals (late Hemingfordian North American Land Mammal Age, represented by Aepycamelus, Copemys and Merychippus; 17.5–15.9 Ma35,41), benthic foraminifera, fossil mollusks, and K/Ar dating all placed the age range between 17.5 and 15 Ma for this geological unit41 in Orange County. More recently, Velez-Juarbe20 revised the age of “Topanga” Formation in this county to 16.5–14.5 Ma based on new foraminiferal zones presented in Ogg et al.42.We propose that USNM PAL V 11367 derives from exposures of the “Topanga” Formation in Riverside County. If this mapped unit in Riverside can be correlated with “Topanga” Formation units in Orange County, it would imply a middle Miocene age, likely 16.5–14.5 Ma20, and given the morphological similarities of this isolated tooth with more complete paleoparadoxiid material in Orange County with stronger age constraints, we think a middle Miocene age for USNM PAL V 11367 is warranted. Given the reduced distribution of outcrops of the “Topanga” Formation24,36 in Corona, we identify two potential localities for USNM PAL V 11367 (Fig. 3). These two localities are situated in urbanized areas, less than 21 km apart, in the northwest and the southeast corners of Corona’s city limits (see Fig. 3B). Both are notably less than 40 km apart from the type locality of N. cecilialina in Orange County, but we urge skepticism for a direct correlation as the marine units of Riverside County requires detailed stratigraphic revision to determine their age constraints; they likely belong to a different depositional basin than “Topanga” Formation exposures in westward Southern California counties.Morphological variation and potential diversity of PaleoparadoxiidaeOur comparisons reveal considerable morphological variation in the arrangement and number of dental cusps across Paleoparadoxiidae (Fig. 4). The cusps arrangement for the m2-3 of Archaeoparadoxia and Paleoparadoxia were previously reported by Inuzuka et al.43 (Fig. 4B), but the addition of another specimen (USNM PAL V 11367) reveals larger morphological variability than previously known for the genus Neoparadoxia (Fig. 4C). Specifically, the holotype of N. cecilialina displays slightly different configurations between its right and left m2, driven mainly by the position of the hypoconulid in occlusal view (Fig. 4C). USNM PAL V 11367, the second known Neoparadoxia m2 (or the first m3), is comparable in size and shape with the same teeth in the type specimen of N. cecilialina, especially the right m2. Both the Smithsonian and LACM specimens display a horizontal alignment of the extra cusp, the hypoconulid, and the entoconid; nevertheless, USNM PAL V 11367 shows a tighter configuration, lacking a wide internal spacing between cusps characteristic of the type specimen of N. cecilialina (Fig. 4C). Given the known ontogenetic changes that affect the dental nomenclature in desmostylians32,44, the addition of more comparative material should help discriminate between competing statements of homology45. The identification of USNM PAL V 11367 from the “Topanga” Formation of Corona represents a second diagnostic record of Neoparadoxia from three separate Middle Miocene units in Southern California, reaffirming its presence as a Middle Miocene taxon: USNM PAL V 11367 from the “Topanga” Formation of Riverside County; Neoparapdoxia (LACM 6920) from the Altamira Shale46; Neoparadoxia from the Topanga Formation of Orange County46,47; and the holotype of N. cecilialina from the lower part of Monterey Formation in the Capistrano syncline, Orange County46. It is possible that other records of Palaeoparadoxiidae from Orange County (e.g.47) and elsewhere in California may represent Neoparadoxia. For example, Awalt et al.32 noted that a palaeoparadoxiid from Orange County identified by Panofsky as Paleoparadoxia sp. (LACM 131889)25 is better referred to Paleoparadoxidae sp., pending a more detailed evaluation of this material, which differs in clear ways from N. ceciliana. One of the benefits of continued descriptive work on desmostylian material from well-constrained stratigraphic contexts in Southern California will be the biostratigraphic opportunities for cross-basin comparisons, especially for exposures of the “Topanga” Formation.Parham et al.46 emphasized that Neoparadoxia occurs widely in middle Miocene units across California: besides the aforementioned ones, Parham et al.46 noted records of this genus from the Sharktooth Hill Bonebed (LACM 120023), the Altamira Shale (LACM 6920), and the Ladera Sandstone15 (UCMP 81302). To date, Neoparadoxia is only known from California, yet it is likely that other paleoparadoxiid material tentatively assigned to other genera may expand the geographic range of this taxon. Interestingly, on the west side of the Pacific (Russia–Japan) and some parts of the east side of the Pacific (Oregon–Washington), Desmostylus spp. and paleoparadoxiids rarely co-occurred from the same formation48,49, yet there are many geological units in South California where desmostylids and paleoparadoxiids co-occurred (e.g., Santa Margarita Formation50,51, Rosarito Beach Formation52, Tortugas Formation51, and Temblor Formation3,4). The abundance of new material from the “Topanga” Formation from Orange and Riverside counties should contribute to the discussion of desmostylian environmental preferences48,53.Lastly, like other marine mammal lineages, desmostylian body sizes reached their maximum body size late in their evolutionary history54. By the middle to late Miocene, desmostylians were the largest herbivorous marine mammals along the North Pacific coastlines54, although they likely competed ecologically with co-occurring sirenians, which later eclipsed desmostylians in body size and survived until historical times in the North Pacific Ocean55. Specifically, in the “Topanga” Formation of Orange County, desmostylians co-occurred with sirenians such as Metaxytherium arctodites56, an ecological association that likely was repeated elsewhere in the mid-Miocene of California (e.g., coeval deposits of the Round Mountain Silt). Given the improving stratigraphic picture of Southern California marine mammal-bearing localities, future work on desmostylian paleoecology could test hypotheses of competition with taxonomic co-occurrence data grounded in strong comparative taphonomic and sedimentological frameworks. More