Status of introduced vertebrates in Galapagos Gustavo Jiménez-Uzcátegui a, Víctor Carrión b, Jabi Zabala a, Paola Buitrón a & Bryan Milstead a

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Status of introduced vertebrates in Galapagos Gustavo Jiménez-Uzcátegui a, Víctor Carrión b, Jabi Zabala a, Paola Buitrón a & Bryan Milstead a a Charles Darwin Foundation, b Galapagos National Park As of May 2007, 36 species of introduced vertebrates have been recorded in Galapagos, 30 of which are now established: fish, 2 amphibians, 4 reptiles, 0 birds, and 3 mammals (Fig. ). The six remaining species were intercepted upon arrival in Galapagos: toad (Bufo sp.) on San Cristóbal in 995; 4 reptiles, including small terrapin (Podocnemis unifilis) on San Cristóbal, Florida turtle (Trachemys scripta), 2 green iguanas (Iguana iguana) on San Cristóbal and Santa Cruz, and five-banded lizard (Eumeces inexpectatus) ready to lay eggs; and mammal, a cotton-head monkey (Saguinus Oedipus) on San Cristóbal, a pet on a private boat (2005-06). Figure. Introduced vertebrate species recorded in Galapagos as of May 2007 Established = 38% Established = 3 Intercepted = 3% 6% Established = Intercepted = 3% Established = 22% Established = 4 Intercepted = 4 Fish Amphibians Reptiles The increase in introduced and established vertebrate species on one or more islands of the archipelago has occurred since human settlers arrived, approximately 50 years ago. However, the conservation and management institutions in Galapagos have achieved considerable success at managing and controlling invasive species at local and regional levels. Control and eradication of large vertebrate species on some islands and islets of the archipelago, as well as the interception upon arrival of six new species in recent years, are examples of achievements in prevention and management (Figs. 2 and 3). Of the 36 species of introduced vertebrates recorded in Galapagos to date, 30 have become established and cause serious damage to the insular ecosystems. 36 Galapagos Report 2006-2007

Figure 2. Number of introduced vertebrate species recorded in Galapagos, by status (established, under control and/or eradicated, and intercepted) Number of Species 35 30 30 Established Controlled and/or eradicated 25 25 Intercepted 9 20 23 20 7 5 2 2 3 0 6 6 5 2 2 5 0 800 825 850 875 900 925 950 975 2000 Note: Year *Species controlled and/or eradicated on one or more islands or islets, not Archipelago-wide. Figure 3. Distribution of introduced vertebrate species (established and eradicated) in the main islands of Galapagos Pinta 3 2 Santiago 3 3 Marchena Bartolomé Genovesa Fernandina 2 Rábida Pinzón 4 22 Santa Cruz Isabela Baltra Puero Ayora Santa Fe 26 San Cristóbal Established introduced vertebrates Eradicated introduced vertebrates Floreana? 8 Española Note: The number of established species does not include those observed or possibly observed in the indicated islands. Galapagos Report 2006-2007 37

Established introduced species Exotic species have been introduced by humans, accidentally or intentionally, since 535. Once established, they cause serious, in some cases irreparable, damage to the natural ecosystems. Therefore, managing, controlling and eradicating introduced species are toppriority goals. Beginning in 968, the Galapagos National Park focused on eradicating goats (Capra hircus), pigs (Sus scrofa), and dogs (Canis familiaris). With advisory and technical support from the Charles Darwin Foundation, control and/or eradication techniques have been applied in specific sites or islands where these species are found. Details on the presence and status of all 36 species of introduced vertebrates, as well as management actions taken on the main islands, are presented in Annex. The historical details on arrival and current status on the different islands or islets of Galapagos are presented below. Fish Tilapia (Oreochromis niloticus) was detected in El Junco lagoon on San Cristóbal Island in 2006 2. An invasive species, it is now regularly monitored and a suitable eradication method is being sought. This species has not been recorded in the other lakes. Frogs The tree frog (Scinax quinquefasciatus) from the coastal zone of mainland Ecuador was recorded for the first time in Galapagos in 998. This species has spread through the lagoons of southern Isabela and has been recorded on Santa Cruz. As of 2002, it had not been recorded on San Cristóbal or Floreana 3. Despite attempts at developing an eradication method, no satisfactory method has been found. Reptiles The first record of introduced geckos was Gonatodes caudiscutatus in 892 and Phyllodactylus tuberculosus in 906 4,5, followed by Lepidodactylus lugubris 6, all in San Cristóbal. On Santa Cruz, P. reissi appeared in 975 6, and later L. lugubris 7. L. Lugubris was also found on Isabela 6. These species compete with endemic species for habitat. So far their management has been limited to research on their distribution and level of competition with endemic species. The smooth-billed ani, Crotophaga ani, was introduced on Isabela in 962 and first recorded on Santa Cruz in 966 and on Santiago in 967 8. It has been recorded on Champion, Gardner-by-Floreana, South Plaza, Marchena, Genovesa, Fernandina (where it is not established but does appear sporadically), Daphne, and Pinta 3. The results of eradication efforts on Marchena are not yet clear. When there is plenty of rainfall, the population increases, as occurred during El Niño in 997 998. The pigeon, Columba livia, was introduced into Galapagos in 972-73 2 and has been recorded in and around populated areas of San Cristóbal, Santa Cruz, and Isabela. It was successfully eradicated from these areas in 2004. The cattle egret, Bubulcus ibis, was recorded in 964 on Santa Cruz, but may have arrived in 960 9. Its nesting was not recorded until 986 0. It is distributed on the four populated islands (Santa Cruz, San Cristóbal, Isabela, and Floreana), mainly in cattle-ranching areas, although it has also been sighted on other islands 8. While it may have arrived by natural migration (there is no available information), it has established itself on the islands because humans have altered the land for ranching and other uses. The Guayaquil red-masked parakeet, Aratinga erythrogenys, was recorded in 996 on San Cristóbal as a pet, not an established population 8. Domestic birds, including chickens (Gallus gallus), ducks (Anas sp.), Guinea hens (Numida meleagris), turkeys (Meleagridis gallipavo), and peacocks (Pavo muticus), were brought to Galapagos by settlers. The quail (Coturnix sp.) was first recorded in 200 3. All are found in urban and rural areas. There are also populations of feral chickens. Tilapia and frogs are both introduced species that were detected in recent years and for which effective eradication methods are still being sought. 38 Galapagos Report 2006-2007

The goat, Capra hircus, had been reported in Galapagos since 685 4. Goats were one of the first introduced species recorded on most islands, and were also the first eradicated from many of them. Goats were recorded on Santiago in 83 5 and 900 6 and on Floreana in 832. 7 Both islands are now monitored to confirm recent eradications. Goats were detected on southern Isabela in 897, and on northern Isabela in 968 8, where they were eradicated in 2006. The first record of goats on Baltra dates back to 900 7 and a few feral individuals remain. Goats were detected on Santa Fe and Española in 905 9 and were eradicated in 97 and 978, respectively 5. On San Cristóbal and Santa Cruz, goats were detected in 847 and 925 7 ; some feral goats remain despite subsistence hunting by Galapagos residents. Goats were first detected on Pinta in 959 20 and then eradicated in 2000 2. The first record of goats on Marchena was in 967 6 and they were initially eradicated in 979 7, and again in 983 3 following an intentional re-introduction. On Rábida, goats were detected in 97 and eradicated that same year 7 and again in 975 3. Their date of introduction to South Plaza is unknown, but they were eradicated in 96 5. Domestic goats are currently found on the four populated islands, with some feral and Judas goats on Isabela and Santiago. Goats were one of the first introduced species to become established in Galapagos and one of the most invasive. The successful eradication of goats from Santiago and northern Isabela was the largest eradication project in the world. Cattle, Bos taurus, were introduced to Floreana in 832, 22 Isabela about 850, Santa Cruz in 923, and San Cristóbal in 84 7. Domestic cattle are currently found on the populated islands. Feral cattle still remain on Isabela and Floreana. The current goal is to eradicate feral cattle from the National Park. Horses, Equus caballus, and donkeys, E. asinus, were introduced to Floreana in 832 5 and 934 23, San Cristóbal in 847, and southern Isabela in 897 5. Records of donkeys on Santiago date back to 875; they were eradicated in 2004 23. At present, donkeys and horses remain in the populated areas (Santa Cruz, San Cristóbal, Isabela, and Florena). There are feral populations on San Cristóbal and Santa Cruz. The date of introduction of sheep, Ovis ariens, on Santa Cruz, San Cristóbal, and Isabela is unknown, but the first record was in 984 7. No wild sheep have been recorded. In 2002, Patry 3 did not record sheep on the main islands, and the species is now considered gone. Guinea pigs, Cavia porcellus, were recorded before 984 in the farming area of Santa Cruz, San Cristóbal, and Isabela. No wild guinea pigs have been recorded 7. In 2002, Patry 3 noted them as disappeared from San Cristóbal, and on Santa Cruz their current status is unknown. Rabbits, Oryctolagus cuniculus, were recorded in the farming areas of Santa Cruz before 989 but no wild rabbits have been recorded 5. Pigs, Sus scrofa, were introduced on Floreana in 832 and then eventually to the other inhabited islands. Pigs were first recorded on San Cristóbal in 835 5 and again in 847 7. On Santiago there were reports of pigs in 875 and again in 878 5 ; they were eradicated from Santiago in 2004 24. The first record on southern Isabela was in 897 and on Santa Cruz about 920 7. At present there are both domestic and feral pigs on the four populated islands (Santa Cruz, San Cristóbal, Isabela, and Floreana). The first cat, Felis catus, was recorded on Floreana in 832. They arrived on Santa Cruz and San Cristóbal with early human settlers 5, and were recorded on Isabela in 869 25. Both feral and domestic cats remain on these islands. They were introduced to Baltra, possibly during World War II, but were eradicated from the island in 2004. The first record of dogs, Canis familiaris, was on Santiago in 685 4, and later on Floreana in 832, San Cristóbal in 842 and 847 7, Isabela in 835 26 or 868 7, and Santa Cruz in 868 7. Feral dogs were eradicated from Floreana and San Cristóbal in 970 26. Domestic dogs are currently found on Santa Cruz, San Cristóbal, Isabela, and Floreana. There are still sporadic reports of feral dogs on Santa Cruz and San Cristóbal. The majority of introduced mammals are also as domestic animals on the four populated islands of the archipelago. Galapagos Report 2006-2007 39

Rodents, including the black rat (Rattus rattus), the Norwegian rat (R. norvegicus), and the house mouse (Mus musculus), arrived in Galapagos with humans, possibly when the islands were first discovered 5. Charles Darwin reported the black rat on Santiago in 835. On Pinzón, it was recorded in 890. Norwegian rats were recorded on Santa Cruz and San Cristóbal in 982. As of 2002, black rats were on the five populated islands (Santa Cruz, San Cristóbal, Isabela, Floreana, and Baltra), Bartolomé, Marielas, Pinzón, and Santiago; Norwegian rats were on the five populated islands and Rábida, and house mice on the five populated islands, Santiago, and South Plaza 3. Research and control programs are being conducted in the National Park, as well as rural and urban areas. Changes in the status of introduced vertebrate species, 999-2006 Significant efforts to eradicate introduced vertebrate species from Galapagos have occurred in recent years, resulting in the successful elimination of several species from some of the major islands and islets. Changes in the status of introduced vertebrate species during the period from 999 to 2006 are indicated in Table 2. Five species have been eradicated from a zone, island, or several islands in the archipelago: cats from Baltra; goats from northern Isabela, Santiago, and Pinta; pigeons from southern Isabela, Santa Cruz, and San Cristóbal; donkeys from northern Isabela and Santiago; and pigs from Santiago. Eradication efforts have resulted in some islands and islets being free of cats, goats, pigeons, donkeys and pigs. Table 2. Changes in the status of introduced vertebrate species on the main islands of Galapagos. STATUS BY YEAR ISLAND Common Name Scientific Name Baltra Cat Felis catus Floreana Quail Coturnix sp. Genovesa Smooth-billed ani Crotophaga ani Northern Isabela Goat Capra hircus Northern Isabela Donkey Equus asinus Southern Isabela Dog Canis familiaris Southern Isabela Pigeon Columbia livia Southern Isabela Quail Coturnix sp. Southern Isabela Donkey Equus asinus Marchena Goat Capra hircus Pinta Goat Capra hircus San Cristóbal Pigeon Columbia livia San Cristóbal Quail Coturnix sp. Santa Cruz Pigeon Columbia livia Santa Cruz Quail Coturnix sp. Santiago Goat Capra hircus Santiago Donkey Equus asinus Santiago Pig Sus scrofa 999 200 2004 2006 Present Disappeared Present Present Present Present Present Eradicated Present Present Present Eradicated Present Present Domestic Domestic Present Present Present Domestic Eradicated Present Present Present Present Eradicated Eradicated Eradicated Present Present Present Eradicated Present Eradicated Eradicated Eradicated 40 Galapagos Report 2006-2007

Annex. Introduced vertebrates in the Galapagos: status, distribution on the main islands, and management actions. Island (or zone) Isabela (North) Isabela (South) Floreana Santa Cruz San Cristóbal Santiago Fernandina Genovesa Rábida Española Pinzón Baltra Pinta Santa Fe Marchena Others (Control) Others (Eradication) Others (Present) Class Common Name Scientific Name Amphibians Tree frog Scinax quinquefasciatus M O O Amphibians Toad Bufo sp. I Guayaquil red-cheeked parakeet Quail Guinea hen Chicken Goose Smooth-billed ani Cattle egret Pigeon Duck Turkey Peacock Donkey Horse Goat Pig Rabbit Guinea pig Cattle Cat Sheep Dog Black rat Norwegian rat House mouse Cotton-headed tamarin Aratinga erithrogenys D Coturnix sp. D? D? D D? Numida meleagris D? D? D D Gallus gallus D D D D D Anser anser D Crotophaga ani P P P P P P O O P O Bubulcus ibis P P P P Columba livia E E E Anas platyrhynchos D D? D D? Meleagris gallopavo D? D? D D? Pavo muticus D? Equus asinus E C UE C C E Equus caballus C D D D Capra hircus E C UE C C E E E E E E E 3 Sus scrofa C UE C C E Oryctolagus cuniculus D? D? Cavia porcellus D? D? D? D? Bos taurus C BE D D Felis catus M M M M M E Ovis aries H H Canis familiaris C C C C Rattus rattus M M M M M P M M O? 6 Rattus norvegicus P? P? M P? P P? Mus musculus P P M M P M 9 Saguinus oedipus I Fish Tilapia Oreochromis niloticus BE Gonatodes caudiscutatus P P Lepidodactylus lugubris P P P Phyllodactylus reissii P Phyllodactylus tuberculosus H Reptiles Green iguana Iguana iguana I I Reptiles Five-banded lizard Eumeces inexpectatus I Reptiles Terrapin Podocnemis unifilis I Reptiles Fresh-water turtle Trachemys scripta? I Legend: E Eradicated, UE Under eradication, C Controlled, D Domesticated, O Observed, I Intercepted, H Historical record, P Present without action, M Mitigation Galapagos Report 2006-2007 4