Zoo Biology 20:545–555 (2001)

DOI 10.1002/zoo.10002

Optimal Conditions for Breeding of

Captive Humboldt Penguins (Spheniscus
humboldti ): A Survey of British Zoos
Nicola Blay1,2† and Isabelle M. Côté1*
1
School of Biological Sciences, University of East Anglia, Norwich, United Kingdom
2
Harewood Bird Garden, Leeds, United Kingdom

We surveyed 16 British zoos and bird gardens to assess the optimal conditions
for breeding of captive Humboldt penguins (Spheniscus humboldti). We obtained
information on population, enclosure, and husbandry characteristics and related
these variables to three measures of per capita breeding success, namely, per
capita egg productivity, chick productivity, and hatching success (measured as
the proportion of eggs laid that hatched). All three fitness measures increased
with an increasing number of breeding pairs and total population size but were
not related to population density. Once the effect of number of breeding pairs
was removed statistically, chick productivity was found to be highest when nest-
ing boxes were lined with sand and gravel instead of alternative substrata such
as twigs or vegetation. Hatching success increased with increasing pool size and
was highest in enclosures with concrete floors. Adult mortality in zoos was gen-
erally low and appeared related to the use of chlorine in freshwater pools and to
the presence of other penguin species in Humboldt penguin displays. Several
enclosure and husbandry parameters were not variable enough to assess their
impact on reproduction of captive Humboldt penguins. Recommendations for
optimising conditions for captive breeding of Humboldt penguins include keep-
ing as large a population as possible in a concrete enclosure with a large pool
area, while providing sand and gravel as nesting material. Bird density may be
important but we did not detect detrimental effects on breeding for densities up
to 0.25 birds m–2. Adult mortality can be minimised by exhibiting Humboldt
penguins in single-species display and avoiding chlorination of pool water. An
experimental approach is recommended to confirm the results of this correla-
tional study. Zoo Biol 20:545–555, 2001. © 2002 Wiley-Liss, Inc.

Key words: sphenicid; husbandry; captive breeding; breeding success

*Correspondence to: Isabelle M. Côté, School of Biological Sciences, University of East Anglia, Nor-
wich, NR4 7TJ, United Kingdom. E-mail: i.cote@uea.ac.uk
†
Present address: International Zoo Veterinary Group, Keighley Business Centre, South Street, Keighley,
West Yorkshire BD21 1AG, United Kingdom.
Received July 7, 2000; Accepted October 5, 2001.

© 2002 Wiley-Liss, Inc.

546 Blay and Côté

INTRODUCTION
Humboldt penguin Spheniscus humboldti is one of the most endangered spe-
cies of penguin [del Hoyo et al., 1993]. It is also one of the species most often
kept in captivity [Cheney, 1989; Edgington, 1990] but least studied in the wild
[Hays, 1986]. The species is native to Peru and Chile, and in the past, breeding
occurred on rocky coastlines and offshore islets [Hays, 1984]. The birds used to
nest on traditional sites, digging burrows out of guano that had accumulated for
years among the pebbles on the rocky shore [Scholten, 1992]. In the early 1800s,
the guano deposits were discovered to be valuable as fertilisers. By the late 1800s,
the collection and sale of guano accounted for 60% of Peru’s gross national prod-
uct [Love, 1994]. The guano deposits on the mainland are now either fully de-
pleted or too shallow to allow Humboldt penguins to excavate nesting burrows
[Love, 1994].
The constant disturbance from guano collectors and destruction of breeding
sites, in addition to the harvesting of penguins and eggs as a source of food
[Crawford and Jahncke, 1999], are thought to be directly responsible for the ini-
tial decline of Humboldt penguins, but recent natural events and human activi-
ties have exacerbated the situation. In the early 1980s, the total Humboldt penguin
population in the wild was estimated at 14,500 to 18,000 individuals [Merritt
and King, 1987]. In 1982–1983, after a strong El Niño Southern Oscillation
(ENSO) event and the collapse of anchovy stocks, the main prey of Humboldt
penguins, breeding failed [Hays, 1986] and population size was reduced to 9,000
birds [del Hoyo et al., 1993]. Since then, non-regulated tourist activities, the
introduction of goats on breeding islands, entanglement of birds in fishing nets,
and another severe El Niño event in 1997–1998 have contributed to the contin-
ued decline of the species in the wild [Culik and Luna-Jorquera, 1997; Simeone
and Schlatter, 1998]. However, for unknown reasons, recovery may be under-
way. Recent censuses of moulting Humboldt penguins during the austral summer
of 2000 yielded a combined population size for Chile and Peru of approximately
30,000 individuals (P. McGill, personal communication).
Humboldt penguins are listed on Appendix I of CITES. The export of birds
from Chile and Peru to zoos around the world ceased in the early 1980s, when
the species was listed as threatened [Boersma, 1991]. Since 1983, no Humboldt
penguin has been legally imported from the wild. Zoos have therefore had to
rely on careful management, including resorting to hand-rearing to increase fledg-
ing success, to maintain their captive populations [Perkins, 1990]. Humboldt pen-
guins are now the subject of several conservation breeding programs, including
the European Endangered Species Programme (EEP) and the American Species
Survival Plan (SSP) organised by the American Zoo and Aquarium Association.
These programs aim to enhance captive populations while maintaining genetic
diversity.
Britain holds a large population of Humboldt penguins, distributed among
several zoological institutions. These zoos present a wide range of husbandry
practices and offer an opportunity to examine quantitatively the success of vari-
ous captive-breeding conditions on Humboldt penguin productivity. The goal of
this study was therefore to collect husbandry and population information about
Humboldt penguins held in British zoos to produce a set of standardised hus-
bandry recommendations.
 Optimal Conditions for Humboldt Penguin Breeding 547

MATERIALS AND METHODS

Survey Information
Population and husbandry data were obtained by contacting 16 British zoos
and institutions known to keep Humboldt penguins and interviewing either the
curator or main keeper responsible for this species. A questionnaire was designed
to standardise the interview. When a visit was not possible, the questionnaire
was sent to the curator or main keeper. We found no significant differences in
the mean values of variables reported to us or recorded on site; hence, we com-
bined the results from all institutions. Seven other institutions with Humboldt
penguin displays did not reply to our request for information.
The questionnaire was broadly divided into two sections, one pertaining to
population data and the other to enclosure and husbandry information. The popu-
lation parameters that were derived from the interviews and questionnaires in-
cluded: 1) population size and sex ratio, 2) number of breeding pairs, 3) egg
productivity (number of eggs per bird per year), 4) chick productivity (number
of chicks surviving for 1 year per bird per year), 5) chick-rearing method (parent
or hand), and 6) population age (i.e., years since the establishment of the captive
population at that institution) and density (number of birds per square meter of
enclosure, including land and water). Egg and chick productivity data were pro-
vided as approximations by keepers of total numbers of eggs and chicks pro-
duced per year. This ensured a higher level of cooperation than requesting more
accurate figures. Information on age structure of each population was also sought,
but too few data were obtained for analysis. Three measures of population fit-
ness were therefore analysed: egg productivity, chick productivity, and hatching
success. The third measure was calculated by dividing chick productivity by egg
productivity and multiplying by 100 to express it as a percentage of eggs laid
that hatched to produce viable chicks (i.e., surviving for at least 1 year). The
husbandry parameters collected included information about the enclosure (dimen-
sion in square meters, substrate type, and type of nesting material provided),
pool (number, area in square meters, water type, and filtration system), level of
disturbance to the birds (frequency of cleaning, location of penguin display within
the zoo [i.e., in a relatively crowded area, e.g., near the entrance or restaurant, or
in more remote area], presence of other species within enclosure, number of cur-
rent keepers, total number of keepers since establishment of population), and
feeding regime (number of times fed per day, food ration). In addition, the num-
ber of birds lost to disease in the past 5 years was noted during the curators/
keepers’ interviews.

Statistical Analysis
Not all zoos could provide all statistics; hence, sample sizes are variable
throughout. Percentages were arcsine/square root transformed for analysis, but
back-transformed means are presented for ease of interpretation. To assess the
effect of enclosure and husbandry parameters on population fitness, we used the
residuals of the regressions between each of the three fitness measures and the
number of breeding pairs in the population, the latter having been shown (see
below) to be a main determinant of population fitness.
548 Blay and Côté

RESULTS
Descriptive Statistics of British Humboldt Penguin Captive Populations
The size of captive Humboldt penguin populations in British zoos varied widely,
from four to 45 birds (mean ± 1 SD = 19.9 ± 12.3 birds). In all institutions, the sex
ratio was close to unity (% males = 50 ± 3.6%), but a sizeable percentage of most
populations remained unsexed (35 ± 29%). There were on average 6.4 (± 4.6) breed-
ing pairs per population (range, 0–15), and captive populations had been established,
on average, for 16.1 ± 14.3 years (range, 1–43).
The three measures of fitness varied greatly among populations, with egg pro-
ductivity ranging from 0 to 1.14 eggs per bird per year, chick productivity from 0 to
0.55 chicks per bird per year, and hatching success from 0% to 92%. There was no
correlation between egg productivity and chick productivity (r = 0.35, n = 15, P =
0.20), nor between egg productivity and hatching success (r = –0.15, n = 12, P =
0.66). However, chick productivity and hatching success were strongly and posi-
tively correlated (r = 0.91, n = 12, P < 0.0001). These two indices may therefore not
be considered completely independent measures of fitness.
Correlates of Fitness: Population Parameters
Population size and the number of pairs in a population appeared to be the best
predictors of population fitness. Egg productivity increased nearly significantly with
the number of breeding pairs (r2 = 0.25, F1,12 = 3.98, P = 0.07; Fig. 1a) and with total
population size (r2 = 0.19, F1,14 = 3.34, P = 0.09). By contrast, egg productivity was
not related to the number of males in the population (r2 = 0.15, F1,14 = 2.05, P =
0.18), the number of females present (r2 = 0.19, F1,12 = 2.88, P = 0.12), population
age (r2 = 0.08, F1,13 = 1.06, P = 0.32), or population density (r2 = 0.07, F1,8 = 0.62, P
= 0.46). Chick-rearing methods had no effect on egg productivity (parent-reared:
0.39 ± 0.34 eggs/bird/yr, n = 9; hand-reared: 0.38 ± 0.28, n = 4; both methods: 0.78
± 0.51, n = 2; F2,12 = 1.17, P = 0.34).
The number of chicks produced per bird per year increased significantly with
the number of breeding pairs (r2 = 0.44, F1,12 = 9.36, P = 0.01; Fig. 1b) and with total
population size (r2 = 0.36, F1,13 = 7.45, P = 0.02), but also with the number of males
(r2 = 0.34, F1,12 = 6.12, P = 0.03) and of females in the population (r2 = 0.38, F1,12 =
7.29, P = 0.02). Chick productivity was related neither to population age (r2 = 0.01,
F1,12 = 0.01, P = 0.99), nor to population density (r2 = 0.01, F1,7 = 0.04, P = 0.85),
and it was not affected by chick-rearing methods (parent-reared: 0.13 ± 0.12 chicks
per bird per year, n = 8; hand-reared: 0.23 ± 0.23, n = 4; both methods: 0.20 ± 0.13,
n = 2; F2,11 = 0.57, P = 0.58).
Hatching success was positively, although not significantly, related to the num-
ber of breeding pairs (r2 = 0.25, F1,10 = 3.35, P = 0.09; Fig. 1c) and to population size
(r2 = 0.27, F1,10 = 3.78, P = 0.08). The proportion of eggs hatched did not change
with the number of males (r2 = 0.23, F1,10 = 2.94, P = 0.12) or of females in the

Fig. 1. Relationship between egg productivity (number of eggs per bird per year) and number of
breeding pairs (regression equation: egg productivity = 0.033 * no. breeding pairs + 0.293) (a); chick
productivity (number of chicks per bird per year) and number of breeding pairs (regression equation:
chick productivity = 0.023 * no. breeding pairs + 0.007) (b); hatching success and number of breeding
pairs (regression equation: hatching success = 0.032 * no. breeding pairs + 0.09) (c).
Optimal Conditions for Humboldt Penguin Breeding 549
550 Blay and Côté

population (r2 = 0.24, F1,10 = 3.11, P = 0.11), with population age (r2 = 0.11, F1,4 =
1.10, P = 0.32) or population density (r2 = 0.34, F1,5 = 2.63, P = 0.17). There was no
difference in hatching success among the different chick rearing methods (parent-
reared: 33 ± 23%, n = 6; hand-reared: 44 ± 46%, n = 3; both methods: 25 ± 0%, n =
2; F2,8 = 0.27, P = 0.77).
Correlates of Fitness: Enclosure and Husbandry Parameters
There was little or no variation in a number of enclosure characteristics. For
example, most populations were held outdoors, had a single pool, which was filled
with freshwater. All but two populations were held in relatively calm parts of the
zoo. All populations were provided with standard nest boxes (i.e., approximately 60
cm high × 60 cm deep × 40 cm wide). The effect of these enclosure characteristics
on population fitness could therefore not be assessed.
None of the enclosure or husbandry parameters measured appeared to affect
egg productivity (all tests, P > 0.10). Similar results were obtained when residual
egg productivity (i.e., controlled for the effect of number of breeding pairs, see Meth-
ods) was examined (all tests, P > 0.10).
The tendency for chick productivity to vary with the type of nesting material
provided (F2,9 = 3.23, P = 0.09) became significant when residual chick productivity
was considered (F2,8 = 7.72, P = 0.01; Fig. 2). The number of chicks produced per
capita, after controlling for the number of breeding pairs in a population, was sig-

Fig. 2. Chick productivity (controlled for number of breeding pairs) on three types of nesting sub-
strata. Means are given ± 1 standard error. Sample sizes are shown above each mean. Samples sizes
with different subscripts indicate that those means are significantly different (Tukey tests, P < 0.05).
 Optimal Conditions for Humboldt Penguin Breeding 551

nificantly greater on sand/gravel than on twigs or a mixture of vegetation and sand

(Tukey tests, P < 0.05 in both cases; Fig. 2). Chick productivity, either uncontrolled
or controlled for breeding pair number, did not vary with any of the other enclosure
or husbandry parameters tested (all tests, P > 0.05).
Hatching success increased significantly with increasing pool size (uncontrolled:
r2 = 0.83, F1,4 = 19.90, P = 0.01, Fig. 3; controlled for number of breeding pairs: r2 =
0.73, F1,4 = 10.99, P = 0.03). The type of substratum present in the enclosure also
influenced significantly hatching success (uncontrolled: F3,8 = 5.06, P = 0.03, Fig. 4;
controlled for number of breeding pairs: F3,8 = 3.86, P = 0.056). Populations held in
concrete enclosures has a significantly higher hatching success than those held on
sand, gravel or a mixture of the two (Tukey tests, P < 0.05, Fig. 4). Hatching success
was not influenced by any of the other enclosure or husbandry variables tested (all
tests, P > 0.05).
Correlates of Recent Adult Losses
The number of adult losses over the past 5 years ranged from 0 to 4 (mean ± 1
SD = 2.0 ± 1.25). The rate of adult losses (number of losses/number of birds in the
population) was significantly higher when Humboldt penguins were displayed with
other penguin species (0.14 ± 0.06 deaths per bird) than in single-species displays
(0.07 ± 0.04 deaths per bird; t = 5.95, df = 9, P = 0.04). Mixed-species displays
usually included African (S. demersus), King (Aptenodytes patagonica), Rockhopper
(Eudyptes crestatus), and/or Gentoo penguin (Pygoscelis papua). The rate of loss

Fig. 3. Relationship between hatching success and pool volume in cubic meters (regression equation:
hatching success = 0.004 * pool volume – 0.024).
552 Blay and Côté

Fig. 4. Hatching success on four types of enclosure substrata. Means are given ± 1 standard error.
Sample sizes are shown above each mean. Samples sizes with different subscripts indicate that those
means are significantly different from each other (Tukey tests, P < 0.05).

was also significantly higher when pool water was chlorinated by the institution (0.16
± 0.06 deaths per bird) than when it was not (0.07 ± 0.04; t = 9.88, df = 9, P = 0.01).
Other population, enclosure, and husbandry variables did not affect recent adult losses
(all tests, P > 0.05).

DISCUSSION
Our survey of British zoos revealed a significant positive effect of increasing
number of breeding pairs and total population size on the population fitness of cap-
tive Humboldt penguins. In addition, we found that a number of enclosure and hus-
bandry characteristics, such as substratum type, nesting material, and pool size,
influenced aspects of reproductive success. Although adult mortality rates were gen-
erally low, higher rates were associated with multi-species displays and chlorination
of pool water.
The effect of breeding pair number and population size on measures of repro-
ductive success, namely, egg and chick productivity, was expected given the social
nature of Humboldt penguin in the wild [Duffy, 1983]. The number of young hatched
per adult has previously been shown to increase with population size for captive
Humboldt penguins [Schmidt, 1978; Olney and Ellis, 1988], as well as captive Afri-
can penguins (S. demersus) and Magellanic penguins (S. magellanicus) [Olney and
Ellis, 1988]. Despite this, population sizes in zoos are typically small (e.g., 20 on
 Optimal Conditions for Humboldt Penguin Breeding 553

average in this study, 15 in Olney and Ellis, 1988). In the wild, it would be unusual
for reproductive individuals to be in contact with fewer than 20 birds [Boersma,
1991]. Our results suggest that increasing the size of a small colony from five to 10
pairs, for example, should more than double per capita chick production.
The importance of breeding pair number or population size was not simply an
artefact of differences in rearing methods in small and large populations. In our study,
the success of parent- versus hand-rearing had no effect on chick productivity or hatching
success. This is in contrast with the results of Cheney [1989] who found that Humboldt
penguin fledging success across a number of zoological institutions was significantly
higher with hand- than with parent-rearing. The success of parent-rearing in Cheney’s
survey (41%) was slightly higher than the figure found in the current study (33%).
Note that Cheney found, as we did, great variation among institutions in success rate
of parent-rearing (range in Cheney, 20–100%; range in our study, 0–69%).
Interestingly, density did not appear to be important to reproductive success.
High densities may have detrimental effects on reproduction because of overcrowd-
ing, competition and spread of disease. Humboldt penguins in British zoos may be
held below the densities at which these detrimental effects would manifest them-
selves. The maximum density reported in our study was 0.25 bird m–2.
Once the effect of breeding pair number on chick productivity was removed,
the importance of nesting material became clear. Sand and gravel were associated
with higher chick productivity than their alternatives, twigs and sand-vegetation mix-
tures. The organic matter in the latter substrates could harbour diseases such as as-
pergillosis, which may affect chick viability. Hatching success, although correlated
with chick productivity, appeared influenced by different factors. High hatching suc-
cess was associated with enclosures with large pools. In fact, pool size explained
nearly 75% of the variation in hatching success among populations. Larger pools
may allow the expression of natural behaviours such as shoaling and porpoising,
resulting in greater well-being of parental birds and hence hatching success. Enclo-
sure substrate type was also important; hatching success was highest in concrete
enclosures. Concrete has been implicated in the development of bumblefoot, i.e.,
abscesses on feet caused by abrasions that affect not only penguins but also birds of
prey and poultry [Siegmund et al., 1967]. Despite this potential problem for adults,
concrete appears to be beneficial to hatching success, perhaps because it offers rela-
tively better hygienic conditions since it is easier to clean than either sand or gravel.
It is important to note that our measure of hatching success does not represent true
hatchability because the data made available to us did not distinguish between hatch-
ing failure due to developmental problems and to infertility. The hatching success
that we calculated for the participating institutions may therefore underestimate true
hatchability, and it is not clear whether pool size and substratum type alleviate either
or both causes of hatching failure.
Overall, adult losses were low, but significantly higher death rates occurred in
mixed-species exhibits. It is possible that contagious diseases are transferred more
readily to Humboldt penguins in such displays, although avian malaria, the highest
cause of mortality in outdoor penguin exhibits, does not usually spread from pen-
guin to penguin [Cranfield et al., 1991]. Alternatively, Humboldt penguins may be at
a competitive disadvantage for food or nest sites if placed with more dominant spe-
cies. Detailed behavioural studies of interspecific interactions are required to deter-
mine the cause of the higher mortality rates in mixed displays, as are examinations
554 Blay and Côté

of post-mortem records to determine whether the causes of mortality differ between

single- and mixed-species displays. More penguins were also lost in exhibits using
chlorinated water. To our knowledge, health problems related to chlorine in penguins
or any other aquatic captive species have not previously been reported.
Several husbandry variables were found to have no effect on egg or chick pro-
ductivity, hatching success, or mortality rate. Among these were the tagging tech-
nique used for individual recognition, pool filtration system, number of keepers, food
ration, and food type. It could be that these factors truly do not affect breeding suc-
cess of Humboldt penguins in captivity, perhaps because zoos have converged to-
wards a number of techniques and practices with proven track records. However, it
is also possible that our sample sizes were too small or that our categorisation was
too crude, in some cases, to detect an effect.
Despite this drawback, we have provided the first quantitative analysis of the
population and husbandry characteristics associated with high reproductive success
in captive Humboldt penguins. Such a quantitative approach is potentially applicable
to a variety of other species that are kept in a number of institutions under different
conditions. Ideally, it should be combined with controlled experimental manipula-
tions, but we believe that a quantitative survey such as ours offers an important first
step in providing recommendations for optimal captive breeding conditions.

CONCLUSIONS
1. Reproductive success of captive Humboldt penguins can be increased by
maintaining as large a population as possible. No detrimental effects of density on
reproductive success were found with densities up to 0.25 bird m–2.
2. Chick productivity can be increased by using sand and gravel as nesting
material instead of twigs or vegetation/sand mixtures.
3. Hatching success of Humboldt penguin eggs is higher in enclosures with
concrete floors and with large pools.
4. To reduce adult mortality, Humboldt penguins should be held in single-spe-
cies displays in enclosures with non-chlorinated pool water. The exact mechanisms
by which mortality is reduced under these circumstances remain to be determined.
5. Experimental manipulations of various enclosure characteristics and hus-
bandry practices should be carried out to confirm these correlational results.

ACKNOWLEDGMENTS
The authors thank all the curators and keepers who made this work possible by
sharing their information and knowledge. The participating institutions included
Birdworld, Chessington Zoo, Chester Zoo, Colchester Zoo, Cotswold Wildlife Park,
Drusillas Park, Dudley Zoo, Flamingoland, Gatwick Zoo, Harewood Bird Garden,
Newquay Zoo, Sewerby Hall, Southport Zoo, Twycross Zoo, Welsh Mountain Zoo,
and Whipsnade Wild Animal Park. Special thanks to Harewood Bird Garden, which
hosted N.B. during the study. We are also grateful to Dr. Patty McGill of Brookfield
Zoo, Illinois, for providing us with the most recent unpublished estimates of wild
Humboldt penguin population size, and to two anonymous reviewers whose com-
ments greatly improved the manuscript. Financial support was provided by the Uni-
versity of East Anglia.
 Optimal Conditions for Humboldt Penguin Breeding
REFERENCES
Boersma PD. 1991. Status of wild and captive pen-
guin populations. TREE 6:381–2.
Cheney C. 1989. Captive reproduction in spheni-
cids: a statistical overview. Sphenicid Penguin
Newslett May:8–9.
Cranfield M, Beall F, Skjoldager M, Ialeggio D.
1991. Avian malaria. Sphenicid Penguin Newslett
Dec:5–7.
Crawford RJM, Jahncke J. 1999. Comparison of
trends in abundance of guano-producing seabirds
in Peru and southern Africa. South Afr J Mar
Sci 21:145–56.
Culik BM, Luna Jorquera G. 1997. Satellite track-
ing of Humboldt penguins (Spheniscus hum-
boldti) in northern Chile. Mar Biol 128:547–56.
del Hoyo J, Elliott A, Sargatal J. 1993. Handbook
of the birds of the world. Vol. 1. Barcelona: Lynx
Edicions.
Duffy DC. 1983. The foraging ecology of Peru-
vian seabirds. Auk 100:800–10.
Edgington DG. 1990. Survey of penguins ex-
hibited in Great Britain and the republic of
Ireland. In: Blackwell S, editor. Penguin man-
agement. Oxford: Cotswold Wildlife Park. p
49–63.
555
Hays C. 1984. The Humboldt penguin in Peru.
Oryx 18:169–80.
Hays C. 1986. Effects of the 1982–1983 El Niño
on Humboldt penguin colonies in Peru. Biol
Conserv 36:169–80.
Love JA. 1994. Penguins. London: Whittet.
Merritt K, King N. 1987. Behavioral sex differ-
ences and activity patterns of captive Humboldt
penguins. Zoo Biol 6:129–38.
Olney PJS, Ellis P. 1988. Int Zoo Yrbk 28:399–400.
Perkins G. 1990. 1989 Regional studbook for
Humboldt penguin. Chicago: Chicago Zoologi-
cal Society.
Schmidt CR. 1978. Humboldt’s penguins at Zurich
Zoo. Int Zoo Yrbk 18:47–52.
Scholten CJ. 1992. Choice of nest-site and mate
in Humboldt penguins (Spheniscus humboldti).
Sphenicid Penguin Newslett Aug:3–13.
Siegmund OH, Eaton LG, et al. 1967. The Merck
veterinary manual: a handbook for diagnosis
and therapy for the veterinarian. Rahway:
Merck and Co.
Simeone A, Schlatter RP. 1998. Threats to a
mixed-species colony of Spheniscus penguins in
southern Chile. Col Waterbirds 21:418–21.