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The Hitchhiker Wave: Non-native Small Terrestrial Vertebrates in the Galapagos

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Abstract

Humans have translocated thousands of species of flora, fauna and microorganisms to places they would never have reached on their own. Non-native species may have effects on biological communities, ecosystem functions and human populations. In island environments, the effects of spreading non-native species on native biodiversity can be severe and lead to native ecosystem transformation and even endemic species extinction. The Galapagos Islands are a region of particular interest and relevance to the issue of species introduction and invasiveness.

In this chapter, I analyse the current status of 25 non-native amphibians, reptiles and birds that have been reported in the Galapagos Islands. Six species have established self-sufficiently in Galapagos and may become invasive: Fowler’s snouted tree frog Scinax quinquefasciatus, common house gecko Hemidactylus frenatus, mourning gecko Lepidodactylus lugubris, dwarf gecko Gonatodes caudiscutatus, Peters’ leaf-toed gecko Phyllodactylus reissii and smooth-billed ani Crotophaga ani. Domestic fowl Gallus gallus holds feral populations, which may have self-sufficient populations, but evidence is unclear. I provide information on the distribution and natural history of non-native species of amphibians, reptiles and birds in Galapagos, including new data about the introduction history of S. quinquefasciatus; evidence on the establishment of H. frenatus on Isabela and San Cristobal islands; the first published record of a non-native snake in Galapagos—Lampropeltis micropholis; the first evidence of predation on squamate reptiles by G. gallus in Galapagos; and evidence of a probable major impact by C. ani due to extensive predation on the endemic Galapagos carpenter bee Xylocopa darwini. I comment on the invasiveness and impact potential of non-native species in Galapagos, identify vulnerable islands for the arrival of non-native species, identify potential hitchhiker that could arrive in the future and propose that it is important to rethink about how we understand, manage and prevent introductions of non-native species. The new wave of introduced species in Galapagos is formed by small hitchhikers, species that are easily overlooked, may travel in high numbers and are highly linked to human-made environments.

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Notes

  1. 1.

    The ecological classification of vegetation is based on the proposal by Wiggins and Porter (1971) .

  2. 2.

    The Global Invasive Species Database (GISD 2010) erroneously reported Eleutherodactylus coqui at Galapagos, citing Snell and Rea (1999) as the source, yet those authors reported Scinax quinquefasciatus .

  3. 3.

    The correct updated name of the toad that arrived to the Galapagos is Rhinella horribilis , assuming its origin was western Ecuador. Until recently, R. horribilis was a synonym of Rhinella marina. However, Acevedo-Rincón et al. (2016) recognised them as different species. Rhinella marina is now restricted to the east of the Andes. Further taxonomic changes are expected, and populations from western Ecuador could receive yet another (new) name (Vallinoto et al. 2010).

  4. 4.

    The non-native populations of Rhinella in Florida have multiple origins, with first individuals coming from Surinam and Colombia. Toads from Surinam were probably Rhinella marina , while those from Colombia could be R. horribilis if their origin was western Colombia or R. marina if they came from eastern Colombia.

  5. 5.

    This name is currently applied to different populations of Scinax that include at least one undescribed cryptic species (R.W. McDiarmid in litt. 2003; S. Ron pers. comm. 2013).

  6. 6.

    Snell and Rea (1999) confused specimens from Isabela with “leptodactylid frogs ”, a common error due to the snout form and general appearance of Scinax frogs.

  7. 7.

    Phillips et al. (2012a) reported a “Tree frog 3 (Hyla sp.)” reported from San Cristobal in 1990. It is possible that it corresponds to early records of Scinax quinquefasciatus . Due to uncertainty with the identification and lack of voucher specimens, they are not included in these analyses.

  8. 8.

    Garman (1892) described Gonatodes collaris , based on two specimens collected by George Baur at Wreck Bay, next to the town of Puerto Baquerizo Moreno, San Cristobal Island. Vanzolini (1965) proposed that G. collaris and G. caudiscutatus were actually synonyms, which was confirmed by Wright (1983) .

  9. 9.

    Several expeditions did not find Gonatodes in San Cristobal Island during the late 1800s and early 1900s (Cope 1889; Heller 1903; Van Denburgh 1912; Slevin 1935). Van Denburgh (1912) , Slevin (1935) and Barbour and Loveridge (1929) suggested that the specimens reported by Garman (1892) were probably collected at Guayaquil, in mainland Ecuador. However, it is probable that G. caudiscutatus was overlooked due to its restricted distribution and low abundance in Puerto Baquerizo Moreno and low activity during the dry season.

  10. 10.

    I observed San Cristobal lava lizard Microlophus bivittatus predating on G. caudiscutatus on June 2005. See account of domestic chicken Gallus gallus for details on a predation event on G. caudiscutatus.

  11. 11.

    All endemic Galapagos geckos which belong to the genus Phyllodactylus are diurnal and nocturnal and inhabit the arid lowlands. They are scansorial and arboreal, having dorsoventrally compressed digits with greatly expanded lamellae. Gonatodes caudiscutatus has a smaller body size than all endemic geckos, is diurnal and mainly inhabits the humid highlands. It is terrestrial and semi-arboreal, having more restricted climbing abilities than the endemic geckos due to its cylindrical digits without expanded lamellae.

  12. 12.

    At least one study on interactions between non-native and endemic geckos in Galapagos has been conducted but remains unpublished (M. Altamirano’s PhD dissertation, cited by Phillips et al. 2012a).

  13. 13.

    Hoogmoed (1989) published the first mention of Lepidodactylus lugubris in Galapagos. However, he did not find the species and cited the unpublished records obtained by John Wright at Puerto Ayora, Santa Cruz Island, in 1983.

  14. 14.

    Marinus Hoogmoed did not find Lepidoblepharis lugubris during his intensive surveys of Puerto Ayora in 1988 (Hoogmoed 1989; Lundh 1998).

  15. 15.

    Until recently, Lampropeltis micropholis was a subspecies of L. triangulum. However, Ruane et al. (2014) raised it to species status. As currently understood, L. micropholis occurs from western Costa Rica to Ecuador. Further taxonomic changes are expected, and populations from the highlands of Ecuador could receive yet another (new) name (J. Valencia, in litt. 2012).

  16. 16.

    Lampropeltis micropholis is rather frequent on the highlands, even in rural and suburban areas. However, there are few specimens from the lowlands (Cisneros-Heredia and Touzet 2007; pers. obs.). Williams (1988) reported it from Guayaquil, based on a specimen collected by Edward Whimper during the 1890s. Pérez-Santos and Moreno (1991) reported the species from the province of Guayas, without providing details. Although no further information about L. micropholis from Guayaquil has been published, I am aware of two additional records: one individual collected ca. 18 km from Guayaquil and delivered to Jean-Marc Touzet (Fundación Herpetológica “Gustavo Orcés” FHGO) in February 1990 (Touzet JM pers. comm.) and another photographed by Keyko Cruz at Cerro Blanco, ca. 8 km from Guayaquil (Cruz 2015).

  17. 17.

    However, Jiménez-Uzcátegui et al. (2007) reported that only one Iguana iguana was found in Santa Cruz in 2000, while the other was found in San Cristobal.

  18. 18.

    Domestic ducks in Galapagos seem to be a mix of descendants from the mallard Anas platyrhynchos and the Muscovy duck Cairina moschata.

  19. 19.

    Japanese quail Coturnix japonica and common quail C. coturnix are distinct but closely related species (Johnsgard 1988; McGowan and Kirwan 2016). Coturnix japonica was domesticated in eastern Asia several centuries ago, and domesticated quails are derived from C. japonica and its hybrids with C. coturnix (Guyomarc’h 2003). While C. coturnix is a partially migratory species, the domestic C. japonica lost its migratory impulse during domestication (Derégnaucourt et al. 2005; Guyomarc’h 2003).

  20. 20.

    Crotophaga ani expansion across America has not been mediated by humans. The species is not listed within the GISD (2010).

  21. 21.

    Still, this introduction hypothesis remains an assumption, mainly based on the apparently low capacity of anis to self-disperse through long distances across oceans (Harris 1973; Grant and Vries (1993), Grant and Grant 1997; Phillips et al. 2012a).

  22. 22.

    When it arrived to Galapagos for the second time, it was weak and died by the next day (Jiménez-Uzcátegui et al. 2015).

  23. 23.

    While all other bird species recorded as vagrants at Galapagos can be classified as oceanic wanderers or as stray boreal migrants (Wiedenfeld 2006; Jiménez-Uzcátegui et al. 2015)

  24. 24.

    The first specimen of Lepidodactylus lugubris from America was collected in Panama in 1916 (Fugler 1966; Hoogmoed and Avila-Pires 2015). G.K. Noble collected it during his trip for the Harvard Peruvian Expedition (Collection catalogue, Herpetology, Museum of Comparative Zoology, Harvard University). The gecko was collected just 2 years after the opening of the Panama Canal and was probably translocated on boats coming from Hawaii or Oceania (Smith and Grant 1961). By 1941, L. lugubris had already reached Colombia (Daza et al. 2012; Hoogmoed and Avila-Pires 2015).

  25. 25.

    In comparison with Santa Cruz Island, the area of urban and suburban gardens in San Cristobal is reduced. This limited habitat availability is apparently the reason why Gonatodes caudiscutatus holds small and restricted populations in the lowlands of San Cristobal and why Scinax quinquefasciatus has not become established in that island (despite its first record in 2000).

  26. 26.

    The largest coastal lagoon of Isabela, Las Diablas, is next to the town of Puerto Villamil. Its low salinity levels (6–10 gL−1, Gelin and Gravez 2002) allow the reproduction of S. quinquefasciatus.

  27. 27.

    Which contributes to their hard detection and improves their survivorship (Olson et al. 2012).

  28. 28.

    Adaptation to desiccation conditions has also enhanced tolerance to salinity in some amphibians (Balinsky 1981; Wells 2007), thus making it easy for them to survive in low salinity lagoons like Las Diablas in Isabela Island. The three species of Scinax that have become established in islands as cargo hitchhikers have adapted to arid environments or anthropogenic areas on their native distributions: Scinax quinquefasciatus, S. x-signatus and S. ruber (Breuil and Ibéné 2008; Breuil 2009; Kraus 2009; Powell et al. 2011). The first two are also known to be adapted to breed in marshes with low salinity (Jiménez-Uzcátegui et al. 2007; Rios-López 2008; pers. obs.). It seems that Scinax species, which are able to adapt to open habitats, show some tolerance to salinity.

  29. 29.

    Frogs that are common in these habitats have easy access to freight or have a great chance to be packed along with horticultural products (Kraus et al. 1999).

  30. 30.

    Species that establish successful self-sufficient populations usually come from areas that have a similar climate to the jurisdiction where they are introduced (Bomford et al. 2009).

  31. 31.

    Eradication attempts by hand capture, spraying caffeine, and increasing the salinity of the lagoons were unsuccessful (Jiménez-Uzcátegui et al. 2007; Phillips et al. 2012b).

  32. 32.

    For example, available knowledge about the populations of Scinax quinquefasciatus in Galapagos remains in two unpublished dissertations: Pazmiño (2011) described the genetic diversity and origin of the Galapagos populations of S. quinquefasciatus, and Vintimilla (2005) analysed the control potential of increasing water salinity.

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Acknowledgements

I am thankful to Gustavo Jimenez-Uzcátegui, Marco Altamirano, Tristan Schramer and Yatin Kalki for providing valuable information about the Galapagos non-native fauna and access to relevant collections; to Marylin Cruz, Arturo Izurieta Valery, Nuria Estrella and David Veintimilla for their kind and prompt response to preserve and study the specimen of Lampropeltis from Galapagos; to Zell Lundberg, Christina Mitchell, Luke Smith, Kevie Dowie and Sandy Espinosa for allowing me to use their photographs; to anonymous reviewers for their useful comments that improved earlier versions of this chapter; to Diego Quiroga, Gabriela Moreno, Cecibel Narváez and Ma. Angélica Moreano for all their help during my stay at Galapagos; and to the Galapagos extension of Universidad San Francisco de Quito USFQ for the logistical support during my field trips to Galapagos Islands in June 2005, June–July 2008 and June–August 2009. The following organisations and projects provided access to literature: King’s College London, Biblioteca Universidad San Francisco de Quito USFQ, Biblioteca Fundación Charles Darwin, Biodiversity Heritage Library (http://www.biodiversitylibrary.org), the Internet Archive (http://www.archive.org), and Google Books (http://books.google.com).

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Cisneros-Heredia, D.F. (2018). The Hitchhiker Wave: Non-native Small Terrestrial Vertebrates in the Galapagos. In: Torres, M., Mena, C. (eds) Understanding Invasive Species in the Galapagos Islands. Social and Ecological Interactions in the Galapagos Islands. Springer, Cham. https://doi.org/10.1007/978-3-319-67177-2_7

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