Galapagos Penguins Swimming Galapagos Penguins Diet The main prey items appear to be thin schooling fish of a length of 1-15 cm, largely around 10 cm long. Sardines (Sardinox sagax), Anchovy (Engraulis sp.), Mullet (Mugil sp.) and Piquitangas (Lile stolifera) are reported prey species. Based on observations of Galapagos Penguins foraging with Audubon Shearwaters, which feed on crustaceans, it is assumed that these may also form part of the diet of the penguins, although this has not been directly proven. Fish are normally swallowed whole, yet penguins have been rarely observed manipulating larger prey items at the surface. Foraging Behaviour Galapagos Penguins are not generally seen porpoising as they leave shore, possibly due to the scarcity of marine predators close to the colonies. However, this behaviour may be observed when they are travelling towards feeding aggregations. Generally underwater swimming involves the movement of the flippers through an arc of about 130'.

During the upstroke the fronts of the flippers are tilted slightly upward to reduce resistance, whilst they are tilted downwards during the downstroke to generate maximum thrust forwards. During surface swimming, the flippers are only moved through a small angle and remain largely submerged. Low water temperatures (below about 23'C) at Galapagos are associated with phases of high marine productivity. Accordingly, during such phases, penguins can be observed in large groups of more than 20 birds, occasionally more than 100. When temperatures are higher, birds are often observed foraging in pairs, scanning the area for small groups of fish. During periods of high productivity, multi-species feeding aggregations may be observed as little as a few meters from the coast. These aggregations, where many penguins may be present, form when large schools of small fish are present. Multispecies feeding aggregations have been studied in detail at Galapagos (Mills 1998. Condor 100, p.277-285). Most occurred less than 150 meters off shore, with about 45% within 50 m.

Galapagos Penguins, together with Brown Pelicans and Brown Noddies were most commonly associated with aggregations close to shore. Penguins tended to dive under the school of fish, picking off fish at the bottom of the school or feeding on the side of the school as they rose to the surface. This behaviour actually appears to fulfil a kind of shepherding role, since aggregations fed on by penguins were longer-lived (often many hours) than those fed on by plunge-divers, which tend to disperse fish. Diving behaviour was studied in two non-breeding male penguin using attached time-depth recorders (Mills 2000. Mar. Ornithol. 28, p.75-79). Travelling and foraging dives were not distinguished, since diving was extremely shallow and no significant transit phase was evident. Mean dive depths of 4 and 2 m, and durations of 25 and 9 sec, were measured for the birds at Fernandina and Bartolome, respectively. Mean distances offshore were 36 and 60 m, respectively, with maxima of only 100 and 300 m observed.

Diving was almost continuous although slightly reduced in frequency around dusk and dawn, when light levels are poorer. At Fernandina, penguins were feeding on Sardines, whilst at Bartolome smaller groups of unidentified fish were being preyed on, often in extremely shallow water. The dive record from the Fernandina penguin showed periods of intense diving to relatively defined depth ranges. Whilst these were not necessarily benthic dives, they presumably were aimed at aggregations of fish at these particular depths. A maximum dive depth of 32 m and duration of over 3 min was reported, showing that Galapagos Penguins generally don't approach their physiological limits when diving. A more extensive study, using GPS and dive loggers to monitor diving behaviour of male and female birds provisioning chicks, was performed in 2004 and 2005 at 3 nesting sites on south-western Isabela (Steinfurth et al. 2008. Endang. Species Res. 4, p.105-112). The penguins moved up to 23.5 km (mean 5.2 km) from their nest sites in foraging trips lasting a mean of 8.4 hours.

Foraging was concentrated in coastal waters less than 500 m from the shore, with several birds approaching a distance of 1 km. As in the previous study, most dives were shallow, with 90% of time spent underwater at depths of less than 6 min. A maximum dive depth of 52 m was however recorded. Male birds tended to travel further than females and mean distances varied from site to site. The reason for the tendency of Galapagos Penguins to forage in close proximity to the coast, which is unique among penguins, was attributed by the authors to either prey preference or avoidance of predation by the numerous shark species around the Galapagos Islands. It is possibly to avoid competition in their limited foraging areas that Galapagos Penguin colonies are generally relatively small. Adult Spheniscus Penguins in general have at least one black ring pattern around their bodies. This may be an adaptation to the pursuit of schooling fish, since one has shown that schools of Cape Anchovies tend to depolarize (break up) more frequently when presented with striped models compared to models without stripes (Wilson et al. 1987. Animal Behav.

35(5), p.1558-1560). Predators are thought to be more efficient at catching individual fish compared to fish in dense aggregations which are formed as a defensive strategy. Hence, the stripe may increase foraging efficiency. Its absence in the other genera of penguins which feed on different types of prey supports this notion. Reproduction: Nest & Partner Selection Galapagos Penguins, especially females, often return to their previous nest sites for breeding. The sites are also frequently visited when no breeding is taking place. Nest burrows are largely found in natural cavities in coastal lava deposits, such as small lava tubes, or may be excavated in tuff, as at Elizabeth Bay. Few poorly-arranged objects such as feathers leaves or twigs line the nest and these are probably ritual gifts rather than functional objects in most cases. It is only necessarily to cushion the eggs when the base of the nest is rough, yet in lava cavities, guano from previous seasons usually smooths the base of the nests. At the onset of a breeding period, unmated birds increasingly spend time visible near the shoreline, possibly in the hope of attracting a mate.

Solitary birds are largely males, since these outnumber female birds. Unmated breeding-age females are rarely observed. Once a pair-bond has been established, it is frequently maintained. In years with base-line levels of mortality, 90% of pairs may be maintained from the previous season, with about 5% divorcing and 5% changing due to loss of a partner. Established pairs tend to spend less time on land prior to egg-laying than newly-formed pairs. Ultimately, it is thought that the female chooses its mate. Body size is not clearly correlated with mate selection. The selection criteria are presently unknown. There are several forms of behaviour associated with courtship. The Galapagos Penguin does however not perform anything resembling an ecstatic display, which is a prominent form of behaviour in many penguin species, including the other Spheniscids. Mutual preening (allopreening) is generally only observed around the breeding season. Standing near to each other, socializing individuals or established pairs initially preen themselves. After a brief pause, one of the birds starts to preen the other on the head or neck. The other bird may continue to preen itself or may reciprocate this behaviour.

Bill duelling is often observed when one bird closely approaches another, sometimes after landing at the beach. The birds face each other and vigorously shake their heads from side to side, resulting in repeated clattering of the tips of the bills. Paired or unpaired penguins may perform this behaviour. In the former case the penguins remain close after performing this ritual and may either simply relax or alternatively flipper-patting and often copulation may follow. Pair interaction (1) Pair interaction (3) Pair interaction (4), just before onset of bill duelling Flipper-patting is performed by the male which rapidly pats his partner as he edges round to the back of her. The male then leans onto the female, gently forcing her to lie prone on the ground. The male climbs onto the back of the female and flipper-patting continues together with some bill interaction as the female penguin raises her bill and both turn their heads from side to side. The male positions itself so as to bring the cloaca in close proximity. For copulation to occur, the female lifts its tale up and the male rounds the back of his body to dip his cloaca onto that of the female.

Cloacal contact and the associated sperm transfer may last for over 30 secs but is usually shorter. After attempted copulation, the male slips off and may walk aside the female often with its head bowed. Both penguins make shake their tails or make swallowing motions. Successful copulation is observed in under 50% of cases and is followed by similar behaviour. Interestingly, copulation attempts have been observed in shallow water by the beach, although it is not known whether such events are productive. Timing of Breeding Galapagos Penguins are the only species without a clearly defined breeding season. Breeding has been observed at any time of year and rather than being seasonally determined appears to occur in response to suitable oceanic conditions, largely correlated with sea-surface temperatures of below about 24'C. Breeding does not commence, or ceases, when water temperatures are higher. Looking at past data, it appears that February-March may be a period with slightly reduced breeding frequency, with May-December being slightly elevated (Boersma 1978. Science 200, p.1481-1483), however these are very weak trends. Breeding may also not be synchronized between colonies at different locations.

Laying and Incubation In the period before egg-laying, Galapagos Penguins spend an increasing amount of time on land. They are often seen standing near the nest site. The males tend to occupy the nests as egg-laying approaches and often are observed braying at dawn and dusk. Both birds generally remain by the nest when the first egg is approaching and the female often remains on the nest during the whole egg-laying period or performs one foraging trip before the second egg arrives 3-4 days after the first. First eggs are longer, whilst second eggs are wider, although there is no significant dimorphism in egg mass. Incubation commences immediately after laying of the first egg. Most penguins only consistently incubate after arrival of the second egg. Desertion rates of eggs are slightly elevated 12-20 days into incubation, although desertion can occur at any time albeit at lower frequency. Incubation Duties Incubation is shared between the pair, with mean lengths of male incubation periods being 1.9 days and of females 2.0 days. During optimal breeding conditions, daily alternation may take place. Absences of up to 10 days have been reported.

Long absences are however often linked to nest desertion. Incubation periods of the eggs are about 38 days. The first egg tends to hatch 2-3 days before the second. The second egg is slightly less likely to hatch, possibly due to lack of fertilization or poorer egg composition. Brood / Guard Phase Generally, both eggs hatch and the chicks are initially guarded for about 25-30 days, before being left alone at the nest site. During the first about 15 days, the chicks are brooded. After this time, the chicks are able to maintain their own body temperatures and the parents presence serves largely to protect against predation by the Sally Lightfoot Crab, Rice Rat or one of the Galapagos Snakes. The guard phase may be slightly longer for single chicks. When conditions are good, provisioning usually occurs once a day as one of the parents returns to the nest in the evening. Observations of the provisioning of one chick revealed that it received about 20 fish of an estimated length of about 10 cm every evening after dark. The female fed more frequently, yet sometimes both mates fed on a single night.

Weight gain by is generally around 35 g per day during productive breeding periods, with older chicks generally gaining weight slightly faster in most cases. Lone chicks gain weight even faster, although the differences are usually only a few grams. When water temperatures are higher, weight gain is generally reduced and differences between younger, older and lone chicks become more pronounced. Commonly, one of the chicks dies, often during the first 2-12 days after hatching, largely of starvation. It is thought that if parental birds are in poor condition they will initially focus on one chick, whilst abandoning the other. If the conditions remain poor, both chicks may be abandoned. Nest desertion rates clearly correlate to higher water temperatures. Adults generally lose weight as chicks grow and present increasing food demands. It has been shown that adults with below-average weights desert their nests first. Desertion is evidently triggered by the need to forage and gain weight to avoid starvation. Where adults were observed to relieve each other at the nest daily, nest desertions were not observed. On the other hand, where one adult was absent from the nest site for 4 or more days on three occasions, nest failure always followed.

Basically, long intervals between nest reliefs indicate food shortage and unwillingness to provision the chicks at the expense of own body condition. Not only the chicks suffer, but also the bird remaining on the nest will lose nearly 50 g per day. This is clearly not tenable over a long time-period. Parents may be observed preening their chicks although this is not reciprocated. Pecking between siblings is common and is usually directed at the smaller one. Creche Phase (Absent) Unlike many other penguin species, including all surface-nesting species, Galapagos Penguin chicks do not form creches. They generally stay in or close to their own nests until shortly before fledging. Accoustic Parent-Chick Recognition Not studied in detail in the Galapagos Penguin. Importance may not be high compared to in some other penguin species, since the nest is the focal point for meeting the chick(s) and no creches form, necessitating recognition of the own chicks. Nevertheless, calls of Spheniscus Penguins are considered individually distinctive, including calls by adults and the so-called peep of the chicks.

Fledging In good breeding seasons, 1.3 chicks per nest can reach fledging age, whereas in poor seasons total failure of nests can occur. By the time they fledge, juveniles have usually attained a body mass approximating to the mean mass of female adult birds. Most body parts approach adult size after 30-40 days, which accounts for the disproportionately large size of e.g. feet and flippers in young penguins. Immature Galapagos Penguins can be recognized by the grey, as opposed to black, feathering on their backs. Further, they lack the body banding of adults and have greyish faces with whitish-coloured cheek patches. The first moult is often performed after about 6 months, after which the adult plumage is obtained. Diffentiation is then only possible based on bill, feet and eye colouration which initially differs from that of older adults. Juvenile Galapagos Penguin(s) Juvenile Galapagos Penguin Adult (left), Juvenile (right) The initial lack of body banding is common to juveniles of all Spheniscus species and may confer a selective disadvantage compared to adults when foraging.

The black band has been shown to increase temporary dispersal of schooling fish, allowing easier predation (Wilson et al. 1987. Animal Behav. 35(5), p.1558-1560). Pre-Breeding Moult Galapagos Penguins moult before breeding. Unpaired males may however moult during the breeding period. The pre-breedng moult is probably an adaptation to the extreme fluctuations in food availability. The moulting period presents the greatest risk of starvation to adult penguins. Hence, it seems that during good conditions the moult is prioritized and performed first, shortly afterwards followed by breeding if conditions remain suitable. Moulting may also occur more than once in certain years. Possibly, avoiding waiting to moult at the last minute when the plumage is severely worn, means that the birds have more flexibility in adapting to changes in conditions. Alternatively, it has been suggested that plumage-wear is high due to the tropical sun and necessitates more frequent moulting. Before moulting, Galapagos Penguins spend about a month foraging to gain weight in advance of the energy-consuming moult process. The mean pre-moult increase in body weight is approx.

400 g, whilst about 600 g are lost during the moult process. Moulting usually involves a period of 15 days on shore. Underweight penguins may return to sea after only 10 days in order to avoid starvation, although the insulation properties of their plumage is probably poor and together with their reduced fat layer they must suffer increased heat loss at sea. Few of these abortive moulters are likely to survive. Female birds are slightly smaller and lighter than males and this may account for their higher mortality during extreme ENSO events, which contributes to the generally higher proportion of males in the population. Penguin (right) has removed most of old feathers and apparently continues to preen. This is relatively unusual. Moulting Galapagos Penguins. Normally old feathers are gradually lost as in right-hand penguin. Moulting Galapagos Penguins The plumage of males and females differs slightly, with male plumage involving slightly bolder markings. Individuals can be recognized by markings on the pink exposed skin at the base of the bill. General Behaviour: The most common and easily recognizable form of maintenance behaviour, preening, is shared by all penguin species.

Galapagos Penguins often spend over an hour preening after returning from sea and may also be observed preening in shallow waters near the coastline. This behaviour serves to clean the plumage and spread the diester wax-rich preen oil exuded by the uropygial (preen) gland to maintain waterproofing. Various stretching motions are interspersed between the preening. In the both-wing stretch, the penguin leans forward, stretches its neck forwards whilst tipping the head upwards, and extends its wings backwards to above the body. The bill may be opened to perform a jaw-stretch. The tail is often wagged after bouts of preening, but also after excretion. Both-Wing Stretch Accompanied by Jaw Stretch Galapagos Penguin Preening Flipper Preening at Sea is Commonly Observed Distributing Preen Gland Excretions on Flipper Aggression and appeasement behaviours are well documented. When one penguin approaches another too closely aggressive behaviour may occur. This involves flattening of the feathers on top of the head, raising of neck feathers and pointing of the bill, possibly with the neck stretched forwards.

It may also involved tipping of the head from side to side, similar to the alternative stare seen in some other penguins. The intruding penguin may return the threats and this may result in "yelling" of the birds at each other and ultimately pecking. On the other hand, the intruder may appease the threatening bird. This apparently involves certain head-movements which are unique, or at least far more pronounced, in Galapagos Penguins, compared to other species. Head-movements involve movement of the bill in an arc. Passing through a slightly lowered central position, the bill is swung about 45' to either side and is lifted to about 140' above the ground at each side of the arc. Penguins perform this behaviour before retreating and are not generally pecked in the process. Juveniles are pecked more than adults since they neither rapidly retreat, nor perform as many head movements as adults. Galapagos Penguins sleep either with their flippers under the body or resting on the side of the body. They do not tuck their bills under their flippers.

Galapagos Penguin resting Galapagos Penguins resting Walking is relatively steady, involving less swaying from side to side than in many other penguins. Tobogganing is rare, due to the unsuitable substrate, and may be only accidental after stumbling when fleeing. Several forms of behaviour associated with mating have been described under the heading "Nest and Partner Selection". Threats: Climate Change / ENSO Events The greatest threat to the survival of the Galapagos Penguin is global warming. Galapagos Penguins are only able to exist around the equator due to the cold Equatorial Undercurrent (Cromwell Current) which flows eastwards through the Galapagos Archipelago and is forced upwards in the vicinity of Fernandina and Isabela Islands. El Nino Southern Oscillation (ENSO) events equally affect the upwelling of the Humboldt Current along the west coast of South America and the upwelling of the Cromwell Current at Galapagos. An increase in upwelling which may last for up to a year and causes high marine productivity generally precedes ENSO events due to strong southeast trade winds increasing circulation of the South Pacific subtropical gyre (Wyrtki 1975. J. Phys.

Oceanogr. 5, p.572-584). However, upwelling is subsequently reduced to below normal levels, leading to high sea surface temperatures in the affected coastal waters and an associated decline in productivity. During intense ENSO events this may lead to severe mortality and breeding failure of seabird populations. The Humboldt Penguin populations in Peru and Northern Chile suffer severely and the entire Galapagos Penguin population is threatened by such events. The magnitude of the problem has been illustrated by past ENSO events. On Fernandina, in 1971, 63 of 82 monitored nests successfully fledged at least one chick. This coincided with productive pre-ENSO conditions. However, during the ENSO event, breeding attempts in Dec. 1972 - Mar. 1973 and Aug. - Oct. 1973 resulted in a total of only one chick from 200 monitored nests (Boersma 1978. Science 200, p.1481-1483).

The weight of adult birds was well below long-term average during the event and several starved adults were found weighing under 1.8 kg, although adult mortality was not extremely high (Boersma 1998. Condor 100, p.245-253). An even more severe ENSO event occurred in 1982-83 and led to huge adult mortality approximating to 77% of the pre-ENSO population (Valle and Coulter 1987. Condor 89, p.276-281). Recovery of the population after this event was slow. This can be attributed to the fact that since the 1980s, there has been a general trend to warmer water temperatures with few and weak El Nina events where water temperatures are lower and productivity is high (Boersma 1998. Condor 100, p.245-253). Thus, the population had not fully recovered when it was hit by a further extreme ENSO event in 1997-98 which reduced the remaining population by about 65%.

Making matters worse, mortality of females was particularly high, leading to an excess of male birds which could not contribute to population recovery (Boersma 1998. Penguin Conserv. Nov. 98, p.10-11). In Humboldt Penguins, nest intrusion behaviour by unmated males resulted in several incidents of egg loss or chick mortality and was responsible for over 10% of breeding failure at a study colony (Taylor et al. 2001. Condor 103(1), p.162-165). If such behaviour also occurs in Galapagos Penguins, this could inhibit population recovery when there is a surplus of male birds. The trends in the Galapagos Penguin population from 1970 to 2003 reveal that recovery following the steep ENSO-induced population declines continued to be slow into the 21st century (Vargas et al. 2005. Ibis 147, p.367-374).

The census data, which is based on marking and resighting techniques involving subsequent extrapolation of the population size based on the proportion of marked birds amongst those birds resighted, can not provide exact figures, yet the population was estimated at 1350 individuals in 2003, compared to 4000 in 1971. The Galapagos Penguin has survived ENSO events for many thousands of years. Several studies have addressed the past frequency of ENSO events by studying lake sediments. For example, the varying salinity of sedimentary strata in hypersaline Bainbridge Crater Lake on the Galapagos Islands was studied (Riedlinger et al. 2002 J. Paleolim. 27, p.1-27). Salinity falls during periods of heavy rainfall which are associated with ENSO events. The study concluded that ENSO events were present but infrequent between 6100 and 4000 yrs BP (before present), but occurred with increasing frequency and intensity in the last 4000 years.

Significant fluctuations in frequency were observed with 152 moderate and 14 strong / very strong events from 2000-1000 BP (before present), but 5 moderate and 36 strong / very strong events in the last 1000 yrs. Another study analysed even older sediments in Laguna Pallcacocha in southern Ecuador (Rodbell et al. 1999. Science 283, p.516-519). Here, sediments were analysed on the basis of layers of material washed into the lake during periods of heavy rainfall. From 15000 to 7000 BP, periods of over 15 years separated periods of heavy rainfall. However, after this period, the frequency of events continually increased, reaching a periodicity of from 2 to 8.5 years by about 5000 BP. After this period, the long-term average frequency was considered similar to that seen at present. The problem is that a relatively rapid increase in ENSO events has been recorded since the late 1960s (Trenberth and Hoar 1997. Geophys. Res. Lett.

34, p.3057-3060) and that there are fewer cool El Nina periods in between during which population levels can recover. Indeed, it appears that equatorial upwelling in general has been decreased by 25% after 1970, with an associated rise in mean sea temperatures of nearly 1'C (McPhaden and Zhang 2002. Nature 415, p.603-608). The survival of the Galapagos Penguin has been modelled based on the continuation of the level and frequency of ENSO events measured in the period between 1964 and 2004 (Vargas et al. 2007. Biol. Conserv. 137, p.138-148). The model estimated a 30% chance of extinction within 100 years. If the frequency of strong ENSO events is doubled, the probability of extinction increases to 80%. Also, the model assumed a 1:1 sex ratio which is at least questionable, especially shortly after severe ENSO events.

If the proportion of females is lower, as suggested by some authors, the outlook is even more bleak. Clearly, any further warming trend will reduce the penguins chances further by increasing mortality of adult and juvenile penguins, and by reducing the number of cool years when the population can recover. Most forecasts are presently pessimistic with regard to the issue of global warming, so it is possible that only captive populations will be able to survive in the foreseeable future (the author is presently not aware of the existence of captive populations). Climatic warming may also be associated with increased wind-speeds and larger waves. Given that most nests are located within 2 m of sea-level, such waves may cause nest flooding. This has indeed been observed in 2004, when at least 4 nests were lost due to a particularly large wave (Vargas et al. 2006. Biol. Conserv. 127, p.107-114). Fishery activities around Galapagos are regulated, yet there is competition between commercial fisheries and penguins for sardines and mullets, which may exacerbate problems during ENSO years.

Further, penguins are known to drown after getting caught in nets deployed to catch Mullet or Sharks. Disease A major potential threat to the population is also considered to be the introduction of disease, and this may exacerbate the population decline resulting from climatic factors. Galapagos Penguins are immunologically relatively naive. Studies have shown that the birds are seronegative for a wide variety of avian viruses, whilst most have antibodies to Chlamydophila psittaci which is generally asymptomatic in penguins. However, occasional outbreaks of highly virulent strains in captive penguins have been reported (Travis et al. 2006. J. Wildlife Dis. 42(3), p.625-632). Further, due to the small population size and associated levels of inbreeding, variation in the genes of the immune system is very low in the MHC loci which are instrumental in disease resistance (Bollmer et al. 2007. Immunogenet. 59, p.593-602).

In general, the Galapagos Penguin has exceedingly low genetic diversity compared to the similar Magellanic or Humboldt Penguins and the population is homogenous in the sense that no subpopulations can be defined based on genetic analysis (Akst et al. 2002. Conserv. Genet. 3, p.375-383; Nims et al. 2008. Conserv. Genet. 9, p.1413-1420). The lack of diversity or subpopulation structure means that the whole population may be ill-equipped if presented with a new pathogen. Further, since the populations at different sites evidently mix, the probability of disease transmission between populations is high. Human settlement and associated introduction of livestock or parasites bear the risks of exposing the penguins to new diseases. The mosquito Culex quinquefasciatus was first reported on the eastern islands of the Galapagos group in 1989 and has since then established itself and spread to other Islands (Whiteman et al. 2005. Ibis 147, p.843-847).

This mosquito has the potential to carry avian malaria and West Nile Virus, in contrast to the native mosquito. Avian Malaria is a major threat to Spheniscus populations in general and can cause over 50% mortality in previously unexposed zoo populations (Cranfield et al. 1991. Sphenisc. Peng. Newslett. 4, p.5-7). It is implicated in the decline of many native forest birds in Hawaii following its introduction (Van Riper 1988. Ecol. Monographs 58, p.111-127). Penguin blood samples taken in 1996 did not reveal evidence for avian malaria or the avian Herpesvirus Marek's Disease Virus (MDV), which had shortly before caused significant mortality in a Galapagos poultry farm (Miller et al. 2000. Mar. Ornithol. 29, p.43-46). MDV may not be able to infect penguins due to its limited host range, yet continued monitoring is required.

Unfortunately, recent research analysing blood samples taken in 2003-2005 has revealed that avian malaria was now already established itself in the Galapagos Penguin population (Levin et al. 2009. Biological Conservation 142, p.3191-3195). Analysis of samples, as in the previous study, involved use of highly sensitive PCR technology which can detect the presence of even minute amounts of Malaria parasite (Plasmodium) DNA. The sequences recovered from the penguins showed little diversity, suggesting that they have recently arrived and had little time to differentiate. Further, they reveal similarity to the species Plasmodium elongatum which is known to cause acute and sometimes fatal infection in penguins. During 2003-2005, levels of Plasmodium infection fluctuated from 3-7% between different sampling periods. There was no obvious upward trend. Infected penguins were detected at various sites on Fernandina, Isabela, Santiago and Bartolome, so essentially the whole range is already affected. No obvious detrimental effects on health were evident and 2 infected birds were recaptured after over half a year.

A major concern is how infection will affect penguins when these are weakened by other factors, such as ENSO events. This remains to be established. Further, the development or arrival of a highly virulent strain is a constant threat. The transmission of disease from poultry is of increasing concern in view of the recent increase in back-yard or large commercial poultry operations on the Galapagos Islands. These provide food for the increasing population and tourist numbers. Seropositivity was found for a number of pathogens in Galapagos chicken stocks during 2001-2003, including Infectious Bursal Disease, Mycoplasma galliseptum, Avian Adenovirus Type I, Marek's Disease, Avian Encephalomyelitis, Infectious Bronchitis Virus and Newcastle Disease Virus (Gottdenker et al. 2005. Biol. Conserv. 126, p.429-439). Many of these are probably endemic in the population, although only few samples seropositive for Newcastle Disease Virus (Avian Paramyxovirus-1) were found, which is a known major threat to penguins.

Disease may pass from poultry to wild bird populations via waste and run-off water from poultry farms but also via interaction between humans or wild animals and poultry that can easily occur in often disease-ridden back yard chicken operations. Pigeons are also regarded as potential transmission sources for avian pathogens and are being actively eradicated (Phillips et al. 2003. Notic. de Galapagos 62, p.6-11). Infections with parasites such as Filarid nematodes are common in the penguin population but are generally considered subclinical or non-pathogenic. Nevertheless, the emergence of new parasites needs to be monitored. Males were preferentially parasitized in the Galapagos Penguins examined, yet no link between parasite load and body mass could be determined (Merkel et al. 2007. J. Parasitol. 93(3), p.495-503). Dirofilaria immitis can probably cause fatality in penguins and is present on Isabela, yet was not detected in the penguins. The chicken populations studied by Gottdenker et al.

also had a number of nematode species and thus could be a source of infections. Predation Predation is not considered to pose a major threat at present. Observations suggest that adults or juveniles suffer occasional predation by Galapagos Hawks (Buteo galapagoensis) on land and by Sharks at sea. It is conceivable that owls could also prey on penguins, as well as Fur Seals, Sea Lions and Orcas, which are known to occasionally prey on other penguin species. Predation by feral house cats and dogs is an increasing problem since human settlements have grown in size in recent years. Rats, Snakes and Crabs may take young or eggs, yet these have generally been deserted beforehand. In fact, only a small percent of even deserted eggs disappear. White-tipped Reef Sharks Sally Lightfoot Crab, most common crab on Galapagos Galapagos Hawk Volcanic Activity Volcanic activity may also locally affect penguin populations. During the 1979 eruption of Cerro Azul on Isabela, lava reached the sea and caused an increase in the water temperature at Bahia Elizabeth.

It has been suggested that this may have negatively affected penguins at this locality (Harcourt 1980. see Valle and Coulter 1987. Condor 89, p.276-281). However, whilst this may cause some disruption, it is unlikely to cause significant fatalities. Recent volcanism has generally been of a diffusive and relatively benign nature on Galapagos and since the Islands have been formed entirely by volcanic processes, the Galapagos Penguin actually owes its existence to these. Impact of Tourism The direct threat caused by tourism is difficult to ascertain. Whilst regulated tourism may have little direct impact on the penguins, rules have often been ignored in the past (De Groot 1983. Biol. Conserv. 26, p.291-300). The situation in this respect seems to be better now. However, the development of touristic infrastructure and need to sustain large numbers of tourists (see e.g. chicken farm issue) has an impact on the Galapagos environment.

Further, the economic benefits of tourism have drawn more people to the islands and the local population continues to rise, with all of the problems generally associated with population growth in environmentally sensitive areas. About 27,000 people live on Galapagos and approx. 100,000 tourists visit annually (Boersma et al., 2005. Science 308, p.925). Many building sites around the periphery of the towns are testimony to the continuing population growth. Where To See: In order to protect the highly sensitive and unique wildlife of the Galapagos Islands, first wildlife reserves were established in 1934 and the whole archipelago, except for some areas of privately owned land, was designated a National Park in 1959. Protection of the wildlife has necessitated strict regulation of visitor activities. Visitors may only visit designated areas and trails in the company of an official naturalist guide. Further, many sites have restrictions on the numbers of visitors. Visitor sites are found on all of the larger islands. Since most sites are only accessible from the sea, tourists generally book day trips by boat or multi-day cruises on small vessels (see e.g.

that travel from island to island. The number of penguins at different sites differs from year to year so it is difficult to suggest where best to view them. Many visitors seem to see small groups around Bartolome, although the less frequented sites on the west coast of Isabela and on Fernandina also offer opportunities. The main nesting sites on Isabela at Caleta Iguana, Playa de los Perros, Las Marielas (Mariela Islands) are not open to visitors, although it is possible to circle the Mariela Islands in a zodiac. There is a small island with a tiny rocky outcrop at the entrance of Puerto Villamil harbour, which frequently has penguins on it, especially at dusk. Puerto Villamil is one of several urban zones in Galapagos and has visitor accommodation. Small boats can be easily organized in the harbour to visit the site although a barrier prevents boats from getting closer than about 10 meters.

Penguin Rock, Puerto Villamil, Isabella, Galapagos Galapagos Penguin, Bartholome Island, by Santiago Island, Galapagos Galapagos Penguin breeding site - Mariela Islands, Isabella Penguins on Mariela Island If lucky enough to encounter the penguins, the guides should make sure that visitors keep at a reasonable distance and do not make any sudden movements or loud noises which may disturb the birds. Nest sites must not be disturbed and moulting birds should be left alone, since this is a critical time for the birds survival. Penguin on rocks at Punta Moreno, Isabella Penguin on rocks at Mariela Island Galapagos Pictures and Photos - Photography - Bird | Wildlife | Nature - Christopher Taylor - Database Search American Oystercatcher [Haematopus palliatus] The American Oystercatcher (Haematopus palliatus), occasionally called the American Pied Oystercatcher, is a member of family Haematopodidae. The bird is uniquely marked by its black and white body and a long, thick orange beak. This shorebird is approxim... - Last revised: Apr 22, 2009 2.

Black-necked Stilt [Himantopus mexicanus] The Black-necked Stilt (Himantopus mexicanus) is a locally abundant resident of American wetlands and coastlines, from the coastal areas of California, much of the interior western United States and along the Gulf of Mexico as far east as Florida, then so... - Last revised: Sep 15, 2013 3. Blue-footed Booby [Sula nebouxii] The Blue-footed Booby (Sula nebouxii) is a bird in the Sulidae family which comprises ten species of long-winged seabirds.The name ï¿½boobyï¿½ comes from the Spanish term bobo, which means \"stupid fellow\". This is because the Blue-footed Booby is clumsy on... - Last revised: Sep 15, 2013 4. Brown Noddy [Anous stolidus] The Brown Noddy or Common Noddy Anous stolidus is a seabird from the tern family.

The largest of the noddies, it can be told from the closely related Black Noddy by its larger size and plumage, which is dark brown rather than black. The Brown Noddy is a t... - Last revised: Apr 22, 2009 5. Brown Pelican [Pelecanus occidentalis] The Brown Pelican (Pelecanus occidentalis) is the smallest of the eight species of pelican, although it is a large bird in nearly every other regard. It is 106-137 cm (42-54 in) in length, weighs from 2.75 to 5.5 kg (6-12 lbs) and has a wingspan from 1.83... - Last revised: Feb 29, 2016 6. The White-vented Storm-petrel (Oceanites gracilis) is a species of seabird in the storm-petrel family Hydrobatidae. The species is also known as Elliot's Storm-petrel. There are two subspecies, O. g. gracilis, which is found in the Humboldt Current off Pe...

- Last revised: Apr 22, 2009 7. Local but fairly common. Two named subspecies occur, mainly in the arid and shore zones: Galï¿½pagos Snake dorsalis (Baltra, Rabida, Santa Cruz, Santa Fe and Santiago), Fernandina Snake occidentalis (Fernandina) and Isabela Snake helleri (Isabela and Tortug... - Last revised: Nov 24, 2016 8. Flightless Cormorant [Phalacrocorax harrisi] The Flightless Cormorant (Phalacrocorax harrisi), also known as the Galï¿½pagos Cormorant, is a cormorant native to the Galï¿½pagos Islands, and an example of the highly unusual fauna there. It is unique in that it is the only cormorant that has lost the abil... - Last revised: Apr 22, 2009 9. The Galï¿½pagos Dove (Zenaida galapagoensis) is a species of bird in the Columbidae family.

It is endemic to the Galï¿½pagos, off Ecuador.Its natural habitats are subtropical or tropical dry forests and subtropical or tropical dry shrubland.... - Last revised: Apr 22, 2009 10. The Large-billed Flycatcher (Myiarchus magnirostris) also known as the Galï¿½pagos Flycatcher is a species of bird in the Tyrannidae family. It is endemic to Ecuador.Its natural habitats are subtropical or tropical dry forests and subtropical or tropical dr... - Last revised: Apr 22, 2009 11. Galï¿½pagos Green Turtle The Galï¿½pagos Green Turtle is a subspecies of the Green Turtle (Chelonia mydas).The Galï¿½pagos Green Turtle is a subpopulation of the Green Sea Turtle, Chelonia mydas. It is endemic to the tropical and subtropical waters of the Pacific Ocean (Koch, 2007).... - Last revised: Aug 06, 2009 12.

Galï¿½pagos Hawk [Buteo galapagoensis] The Galï¿½pagos Hawk, (Buteo galapagoensis), is a large hawk endemic to the Galï¿½pagos Islands. Known for its fearlessness towards humans and authority over the islands as the only original predator, this raptor has inhabited the Galï¿½pagos archipelago for ov... - Last revised: Apr 22, 2009 13. Galï¿½pagos Mockingbird [Nesomimus parvulus] The Galï¿½pagos Mockingbird (Nesomimus parvulus) is a species of bird in the Mimidae family. It is endemic to the Galï¿½pagos Islands, Ecuador.The Galï¿½pagos Mockingbird is easily spotted on the Galï¿½pagos Islands due to its feathers which are streaked brown an... - Last revised: Apr 22, 2009 14.

Galï¿½pagos Penguin [Spheniscus mendiculus] The Galï¿½pagos Penguin (Spheniscus mendiculus) is a penguin endemic to the Galï¿½pagos Islands. It is the only penguin to live on the equator and can survive due to the cool temperatures resulting from the Humboldt Current and cool waters from great depths b... - Last revised: Apr 22, 2009 15. Galï¿½pagos Sea Lion The Galï¿½pagos Sea Lion (Zalophus californianus wollebaeki) is a species of mammal in the Otariidae family endemic to Ecuador. Being fairly social, and one of the most numerous of the archipelago species, they are often spotted sun-bathing on sandy shores... - Last revised: Apr 19, 2009 16. Galï¿½pagos Shearwater [Puffinus subalaris] The Galï¿½pagos Shearwater, Puffinus subalaris is a small shearwater.

Until recently it was considered to be a subspecies of Audubon's Shearwater, but it is actually one of two members of a very ancient lineage of the small Puffinus species, the other being... - Last revised: Apr 22, 2009 17. Galï¿½pagos Tortoise The Galï¿½pagos tortoise (or Galï¿½pagos giant tortoise), is the largest living tortoise, endemic to nine islands of the Galï¿½pagos archipelago. Adults of large subspecies can weigh over 300 kilograms (660lb) and measure 1.2 meters (4 ft) long. Although the ma... - Last revised: Aug 06, 2009 18. Galï¿½pagos Yellow Warbler [Dendroica petechia aureola] The Yellow Warbler, Dendroica petechia, is a New World warbler. It is the most widespread Dendroica warbler, breeding in almost the whole of North America and down to northern South America.The Yellow Warbler proper (aestiva group) breeds in the whole of...

- Last revised: Apr 22, 2009 19. Great Frigatebird [Fregata minor] The Great Frigatebird (Fregata minor) is a large dispersive seabird in the frigatebird family. Major nesting populations are found in the Pacific (including Galï¿½pagos Islands) and Indian Oceans, as well as a population in the South Atlantic. It feeds on f... - Last revised: Apr 22, 2009 20. Hood Mockingbird [Nesomimus macdonaldi] The Hood Mockingbird (Nesomimus macdonaldi) or Espaï¿½ola Mockingbird is a species of bird in the Mimidae family. It is endemic to Espaï¿½ola Island in the Galï¿½pagos Islands, Ecuador and is a member of the Galï¿½pagos mockingbirds, four closely related species... - Last revised: Apr 22, 2009 21.

Land Iguana [Conolophus subcristatus] The Galï¿½pagos Land Iguana (Conolophus subcristatus) is a species of lizard in the Iguanidae family. It is one of two species of the genus Conolophus. It is endemic to the Galï¿½pagos Islands, primarily the islands of Fernandina, Isabela, Santa Cruz, North S... - Last revised: Aug 06, 2009 22. Large Cactus-Finch [Geospiza conirostris] The Large Cactus-finch (Geospiza conirostris) is a species of bird often placed in the Emberizidae family. It is one of Darwin's finches, and is endemic to the Galï¿½pagos islands (Ecuador), breeding only on Espaï¿½ola Island, Genovesa, Darwin Island, and Wol... - Last revised: Dec 07, 2011 23. Large Ground-finch [Geospiza magnirostris] The Large Ground-finch (Geospiza magnirostris) is a species of bird.

One of Darwin's finches, it is now placed in the family Thraupidae and was formerly in the Emberizidae. It is endemic to Ecuador. Its natural habitats are subtropical or tropical dry for... - Last revised: Apr 22, 2009 24. Lava Gull [Larus fuliginosus] The Lava Gull (Larus fuliginosus) is a large gull and very closely related to the Laughing Gull. It is the world's rarest gull with the entire population living on the Galï¿½pagos Islands. There is only an estimated 300 pairs there.Adult plumage, acquired i... - Last revised: Apr 22, 2009 25. Lava Heron [Butorides sundevalli] The Lava Heron, Butorides sundevalli, is a species of heron endemic to the Galï¿½pagos Islands. The adult is slate-grey, which helps it blend in with the hardened lava. The back feathers typically have a silvery sheen and it has a short crest on its head. W...

- Last revised: Apr 22, 2009 26. The reptile genus Tropidurus includes several species of ground lizard. It includes seven which are endemic to the Galï¿½pagos Islands, where collectively known as lava lizards, although these commonly are placed in the genus Microlophus instead. The distri... - Last revised: Aug 06, 2009 27. Magnificent Frigatebird [Fregata magnificens] The Magnificent Frigatebird (Fregata magnificens) was sometimes previously known as Man O'War, reflecting its rakish lines, speed, and aerial piracy of other birds.It is widespread in the tropical Atlantic, breeding colonially in trees in Florida, the Car... - Last revised: Feb 04, 2013 28. Marine Iguana The Marine Iguana (Amblyrhynchus cristatus) is an iguana found only on the Galï¿½pagos Islands that has the ability, unique among modern lizards, to live and forage in the sea. It has spread to all the islands in the archipelago, and is sometimes called the...

- Last revised: Oct 22, 2011 29. Medium Ground-finch [Geospiza fortis] The Medium Ground-finch (Geospiza fortis) is a species of bird in the Emberizidae family.It is endemic to Ecuador. Its natural habitats are subtropical or tropical dry forests and subtropical or tropical dry shrubland.One of Darwin's finches, the species... - Last revised: Apr 22, 2009 30. Nazca Booby [Sula granti] The Nazca Booby, Sula granti, is a booby which is found in the eastern Pacific Ocean, namely on the Galï¿½pagos Islands where it can be seen by eco-tourists, and on Clipperton Island. The Revillagigedo Islands off Baja California which possibly constitute i... - Last revised: Aug 17, 2009 31.

Red-billed Tropicbird [Phaethon aethereus] The Red-billed Tropicbird, Phaethon aethereus, also known as the Boatswain Bird is a tropicbird, one of three closely related seabirds of tropical oceans. It occurs in the tropical Atlantic, eastern Pacific and Indian Oceans.The Indian Ocean race of the R... - Last revised: Apr 22, 2009 32. Red-footed Booby [Sula sula] The Red-footed Booby, Sula sula, is a large seabird of the gannet family, Sulidae. They are powerful and agile fliers, but they are clumsy in takeoffs and landings.It's the smallest of all boobies, at 71 cm in length and a 137 cm wingspan. It has red legs... - Last revised: Apr 22, 2009 33. Ruddy Turnstone [Arenaria interpres] The Ruddy Turnstone (Arenaria interpres) is a small wading bird, one of two species of turnstone in the genus Arenaria.

It is now classified in the sandpiper family Scolopacidae but was formerly sometimes placed in the plover family Charadriidae. It is a... - Last revised: Oct 28, 2015 34. Sally Lightfoot Crab The crab Grapsus grapsus (known variously as \"red rock crab\", \"abuete negro\", and, together with other crabs such as Percnon gibbesi, as \"Sally Lightfoot\") is one of the most common crabs along the western coast of South America. It can also be seen... - Last revised: Aug 06, 2009 35. Small Ground-finch [Geospiza fuliginosa] The Small Ground-finch (Geospiza fuliginosa) is a species of bird in the Emberizidae family. It is endemic to Ecuador. Its natural habitats are subtropical or tropical dry forests and subtropical or tropical dry shrubland.... - Last revised: Apr 22, 2009 36.

Smooth-billed Ani [Crotophaga ani] The Smooth-billed Ani Crotophaga ani is a large near-passerine bird in the cuckoo family. It is a resident breeding species from southern Florida, the West Indies, Costa Rica, Trinidad and Tobago, south to western Ecuador, Brazil ,northern Argentina and t... - Last revised: Sep 29, 2011 37. Swallow-tailed Gull [Creagrus furcatus] The Swallow-tailed Gull (Creagrus furcatus) is an equatorial seabird in the gull family Laridae. The species is endemic to the Galï¿½pagos Islands. When it is not breeding it is totally pelagic (flying and hunting over the open oceans), migrating eastward t... - Last revised: Apr 22, 2009 38. Warbler Finch [Certhidea olivacea] The Warbler Finch (Certhidea olivacea) is a species of bird in the Emberizidae family.

It is the only member of the genus Certhidea.It is endemic to Ecuador.Its natural habitats are subtropical or tropical dry forests, subtropical or tropical moist montan... - Last revised: Apr 22, 2009 39. Waved Albatross [Phoebastria irrorata] The Waved Albatross, Phoebastria irrorata - also known as Galï¿½pagos Albatross - is the only member of the Diomedeidae family located in the tropics. When they forage, the Waved Albatross follow straight paths to a single site off the coast of Peru, about... - Last revised: Apr 22, 2009 40. Wedge-rumped Storm-Petrel [Oceanodroma tethys] The Wedge-rumped Storm-petrel, Oceanodroma tethys, is a storm-petrel.Common resident to the Galï¿½pagos Islands and endemic subspecies tethys. Population estimated at c. 200,000 pairs in 3 colonies. Nests colonially in burrows or crevices, breeding througho...

- Last revised: Apr 22, 2009 41. Whimbrel [Numenius phaeopus] The Whimbrel Numenius phaeopus, is a wader in the large family Scolopacidae. It is the one of the most widespread of the curlews, breeding across much of subarctic North America, Europe and Asia as far south as Scotland.This is a migratory species winteri... - Last revised: Sep 29, 2011 42. White-cheeked Pintail [Anas bahamensis] The White cheeked Pintail or Bahama Pintail (Anas bahamensis) is a dabbling duck of the Caribbean, South America and the Galï¿½pagos Islands.This species was first described by Linnaeus in his Systema naturae in 1758 under its current scientific name. There... - Last revised: Sep 29, 2011 43.

White-vented Storm-Petrel [Oceanites gracilis] The White-vented Storm-petrel (Oceanites gracilis) is a species of seabird in the storm-petrel family Hydrobatidae. The species is also known as Elliot's Storm-petrel. There are two subspecies, O. g. gracilis, which is found in the Humboldt Current off Pe... - Last revised: Apr 22, 2009 44. Woodpecker Finch [Camarhynchus pallidus] The Woodpecker Finch, Camarhynchus pallidus, is one of the so-called Darwin's finches. First described by Charles Darwin on the Galï¿½pagos Islands, it is a unique species which uses a twig, stick, or cactus spine as a tool. The tool is used as compensation... - Last revised: Apr 22, 2009 45.

Question: Spheniscus mendiculusis the scientific name for the only Penguin to live in equatorial waters; by what name is it commonly known?
Answer: GALAPAGOS PENGUIN