2. WADERS,DIKKOPS, COURSERS AND COMMON LARKS AT OUDTSHOORN

Eggleton, P. &; Siegfried, W. R. 1979. Displays of the Jackass Penguin. Ostrich 50:139-167.

The paper describes the displays of the Jackass Penguin Spheniscus demersus. It also covers aspects of the species' comfort behaviour. Observations were made at several islands off the South African west coast from 1973 to 1976, although most were made on 130 individually-marked adults at Bird Island (Lambert's Bay) in 1975.

The Jackass Penguin has an extensive repertoire of aggressive displays including Point, Gape, Alternate Stare, Sideways Stare, Peck and Beak Slapping. Other displays are Vibratory Head Shaking (VHS), Ecstatic Display, Mutual Ecstatic Display, several bowing displays and displays associated with copulation. The functions of the displays are discussed and comparison is made with displays in other penguins as documented in the literature. In general the displays of the Jackass Penguin resemble those of other penguins, excluding the genus Aptenodytes. There are, however, variations commensurate with the environment and habits of S. demersus.     

Age‐specific survival and movement among major African Penguin Spheniscus demersus colonies

Reliable estimates of survival and dispersal are crucial to understanding population dynamics, but for seabirds, in which some individuals spend years away from land, mortality and emigration are often confounded. Multistate mark–recapture methods reduce bias by incorporating movement into the process of estimating survival. We used a multistate model to provide unbiased age‐specific survival and movement probabilities for the Endangered African Penguin Spheniscus demersus based on 5281 nestlings and 31 049 adults flipper‐banded and resighted in the Western Cape, South Africa, between 1994 and 2012. Adult survival was initially high (≥ 0.74) but declined after 2003–2004 coincident with a reduction in the availability of Sardine Sardinops sagax and Anchovy Engraulis encrasicolus on the west coast of South Africa. Juvenile survival was poorly estimated, but was lower and more variable than adult survival. Fidelity to the locality of origin varied over time, but was high in adults at Robben and Dassen islands (≥ 0.88) and above 0.55 for juvenile and immature Penguins at all localities. Movement occurred predominantly during 1994–2003 and was indicative of immigration to Robben and Dassen islands. Our results confirm that a prolonged period of adult mortality contributed to the observed decline in the African Penguin population and suggest a need for approaches operating over large spatial scales to ensure food security for marine top predators.     

Modularity and Complete Natural Homeoses in Cervical Vertebrae of Extant and Extinct Penguins (Aves: Sphenisciformes)

The cervical system of extant penguins (Aves: Sphenisciformes) is organised into morphological modules, each with its biomechanical function. Indeed, for these marine pelagic birds to acquire hydrodynamic morphology, the folding of the neck is essential. Despite a common general structure, the cervical vertebrae exhibit morphological differences depending on their positioning. These characteristics are identified as apparent cases of complete natural homeotic transformations—therefore, the composition of some modules varies. Two types of complete cervical homeoses are identified between species, but the second type can also occur within some species when the post hatching development is considered. The fossil material analysed here makes it apparent that the two modular configurations characterising the anterior part of the neck—a consequence of the first homeosis—existed 36 My and 25 My ago, for one, and circa 10 My ago, for the other. These comparisons also reveal a clear differentiation in vertebral features between the fossil species of the Oligocene–Miocene ages and the more recent and extant penguins. Ultimately, these observations make the proposal of a hypothesis in relation to the ontogenetic influence of Hox genes, and their regulators, based on the changes observed in the cervical segment of Sphenisciformes.     

In stark contrast to widespread declines along the Scotia Arc,a survey of the South Sandwich Islands finds a robust seabird community

The South Sandwich Islands, in the South Atlantic Ocean, are a major biological hot spot for penguins and other seabirds, but their remoteness and challenging coastlines preclude regular biological censuses. Here we report on an extensive survey of the South Sandwich Islands, the first since the late 1990s, which was completed through a combination of direct counting, GPS mapping, and interpretation of high-resolution commercial satellite imagery. We find that the South Sandwich Islands host nearly half of the world’s Chinstrap Penguin (Pygoscelis antarctica) population (1.3 million breeding pairs), as well as c. 95,000 breeding pairs of Macaroni Penguins (Eudyptes chrysolophus), and several thousand breeding pairs of Gentoo Penguins (Pygoscelis papua). Despite being at the northern edge of their breeding range, we found an unexpectedly large (≥125,000 breeding pairs) population of Adélie Penguins (Pygoscelis adeliae). Additionally, we report that nearly 1900 pairs of Southern Giant Petrels (Macronectes giganteus) breed in the South Sandwich Islands, 4 % of the global population, almost all of which are found on Candlemas Island. We find that the South Sandwich Islands have not experienced the same changes in penguin abundance and distribution as the rest of the Scotia Arc and associated portions of the western Antarctic Peninsula. This discovery adds important context to the larger conversation regarding changes to penguin populations in the Southern Ocean.     

Assessment of aphid colonies by hoverflies. II pea aphids and 3 syrphid species;Betasyrphus serarius (Wiedemann),Metasyrphus frequens Matsumura andSyrphus vitripennis (Meigen) (Diptera: Syrphidae)

In the aphidophagous syrphid species,Betasyrphus serarius (Wiedemann),Metasyrphus frequens Matsumura andSyrphus vitripennis (Meigen), females in search of oviposition sites assess the qualitative and quantitative value of pea aphid colonies for securing the successful development of their offspring. They select, as their oviposition sites, young and “promising” aphid colonies consisting of nymphs and/or adult aphids including few winged adults. They neglect large, older colonies of 4th-instar nymphs and/or winged adult aphids. The 3 syrphid species adopt “buy-futures” ovipositional tactic as doesEpisyrphus balteatus (de Geer), although some tinge of “spot-transaction” is noticed inM. frequens andS. vitripennis.     

Short-term responses of king penguins Aptenodytes patagonicus to helicopter disturbance at South Georgia

The short-term behavioural effects of helicopter overflights on breeding king penguins Aptenodytes patagonicus at South Georgia were examined. Seventeen helicopter overflights were made at altitudes between 230 and 1,768 m (750–5,800 ft) above ground level. Noise from the aircraft engines and helicopter blades increased sound levels in the colony from a background level of 65–69 dB(A) to a maximum mean peak level of 80 dB(A) during overflights. Penguin behaviour changed significantly during all overflights at all altitudes compared to the pre- and post-flight periods. Pre-overflight behaviour resumed within 15 min of the aircraft passing overhead and no chicks or eggs were observed to be taken by predators during overflights. Non-incubating birds showed an increased response with reduced overflight altitude, but this was not observed in incubating birds. Variability in overflight noise levels did not affect significantly the behaviour of incubating or non-incubating birds. Penguins exhibited a reduced response to overflights as the study progressed (despite later flights generally being flown at lower altitudes) suggesting some degree of habituation to aircraft. To minimise disturbance to king penguins we recommend a precautionary approach such that overflights are undertaken at the maximum altitude that is operationally practical, or preferably are avoided altogether.     

The functions of status in the mantled howler monkey,Alouatta palliata Gray: Intraspecific competition for group membership in a folivorous neotropical primate

In the dominance hierarchies of adult male and female mantled howler monkeys (Alouatta palliata Gray), high-ranking individuals are young adults; intermediate-ranking individuals, middle-aged adults; and low-ranking individuals, old adults. This relationship reverses the trend observed in most group-living animals and is previously unreported for this species. A limiting supply of palatable leaves may create intense intraspecific competition for group membership which, it is hypothesized, has resulted in this rare pattern of hierarchical relations. It is shown that individual-level selection is sufficient to explain the evolution of this apparently “altruistic” status system, though other mechanisms are assessed. The energetic constraints imposed by a folivorous diet appear to restrict the expression of aggression to “ritualized” forms. Two groups of monkeys in two different habitats were studied and the rates of appeasement and aggressive behavior were found to be higher in the more “stressful” forest. The latter group, however, displays significantly more behavioral patterns entailing low or intermediate energy expenditure.     

The use of stable isotope ratios in whitebait otolith carbonate to identify the source of prey for Western Australian penguins

A methodology has been developed and used to investigate a direct link between juvenile whitebait (Hyperlophus vittatus), the key prey species sampled from the stomachs of Little Penguins (Eudyptula minor), and the specific nursery area of that prey. A unique application of existing methodology initially involved the measurement of the stable isotopes of oxygen (δ¹⁸O:δ¹⁶O) and carbon (δ¹³C:δ¹²C) in sagittal otolith carbonate, using standard mass spectrometric techniques. The results indicated that the inshore distribution of juvenile (0+ year old) whitebait between Cockburn Sound and Koombana Bay, Western Australia consisted of a number of separate assemblages. Measured differences in stable oxygen isotope ratios were attributed to variations in freshwater input to the embayments that provided the whitebait habitats. In contrast, the measured stable carbon isotope ratios probably resulted from the different isotopic compositions of the food webs in the various habitats. Secondly, a comparison of the average value of carbon and oxygen isotope signatures of pooled otoliths from samples of whitebait from a number of different nearshore coastal sites (assemblages), with that of whitebait obtained from the stomachs of penguins at their main breeding site (Penguin Island) indicated that the values from the penguins resemble most closely those of the average otolith values obtained from whitebait from only one site (Becher Point). Assuming that the whitebait sampled were representative of the whitebait in the nearshore habitats and the diets of the penguins, then these results imply that at the time of sampling the penguins were feeding on whitebait from only one site.     

Detection, differentiation, and abundance estimation of penguin species by high-resolution satellite imagery

Due to its high spatial resolution, broad spatial coverage, and cost-effectiveness, commercial satellite imagery is rapidly becoming a key component of biological monitoring in the Antarctic. While considerable success in surveying emperor penguins (Aptenodytes forsteri) has been facilitated by their large size and the visual simplicity of their habitat, there has been considerably less progress in mapping colonies on the Antarctic Peninsula and associated sub-Antarctic islands where smaller penguin species breed on topographically complex terrain composed of mixed substrates. Here, we demonstrate that Adélie penguin (Pygoscelis adeliae), chinstrap penguin (P. antarcticus), gentoo penguin (P. papua), and macaroni penguin (Eudyptes chrysolophus) colonies can be detected by high-resolution (2-m multispectral, 40–50-cm panchromatic) satellite imagery and that under ideal conditions, such imagery is capable of distinguishing among groups of species where they breed contiguously. To demonstrate the potential for satellite imagery to estimate penguin population abundance, we use satellite imagery of Paulet Island (63°35′S, 55°47′W) to estimate a site-wide population of 115,673 (99,222–127,203) breeding pairs of Adélie penguins.     

Penguins, fur seals, and fishing: prey requirements and potential competition in the South Shetland Islands, Antarctica

Antarctic and sub-Antarctic seabirds, marine mammals, and human fisheries concentrate their foraging efforts on a single species, Antarctic krill (Euphausiasuperba). Because these predators may have a significant effect on krill abundance, we estimated the energy and prey requirements of Adelie (Pygoscelisadeliae), chinstrap (Pygoscelisantarctica), and gentoo (Pygoscelispapua) penguins and female Antarctic fur seals (Arctocephalusgazella) breeding on the South Shetland Islands, Antarctica and compared these estimates with catch statistics from the Antarctic krill fishery. Published data on field metabolic rate, population size, diet, prey energy content, and metabolic efficiency were used to estimate prey requirements of these breeding, adult, land-based predators and their dependent offspring. Due to their large population size, chinstrap penguins were the most significant krill predators during the period examined, consuming an estimated 7.8 × 10⁸ kg krill, followed by Adelie penguins (3.1 × 10⁷ kg), gentoo penguins (1.2 × 10⁷ kg), and Antarctic fur seals (3.6 × 10⁶ kg). Total consumption of all land-based predators on the South Shetland Islands was estimated at 8.3 × 10⁸ kg krill. The commercial krill fishery harvest in the South Shetland Island region (1.0 × 10⁸ kg) was approximately 12% of this. Commercial harvest coincides seasonally and spatially with peak penguin and fur seal prey demands, and may affect prey availability to penguins and fur seals. This differs from the conclusions of Ichii et al. who asserted that the potential for competition between South Shetland predators and the commercial krill fishery is low. Received: 26 August 1997 / Accepted: 16 December 1997     

A revision of<Emphasis Type="Italic">Poposaurus gracilis</Emphasis> (Archosauria: Suchia) based on two new specimens from the Late Triassic of the southwestern U.S.A.

Two associated partial skeletons, one from the McCarty ranch (MOTTU site 0690) near Kalgary, Texas, Tecovas Formation, and the other from the Petrified Forest Member of the Chinle Formation, outside the Petrified Forest, Arizona, represent the most complete specimens ofPoposaurus gracilis found to date, and increase our understanding of the osteology of this rare pseudosuchian archosaur. The cervical centrum of TTUP-10419 possesses an accessory dorsal parapophyseal rib facet, as in “Lythrosuchus”. The ilia of these new fossils are the first ofPoposaurus to preserve the complete preacetabular blade, which is long and “blade-like” rather than “paddle-shaped” as previously thought. These specimens also confirm that isolated elements previously referred toPoposaurus (fused sacral vertebrae, the pubes and the ischia), but not present in the holotype indeed belong to that taxon. The morphology of the cervical centrum and the preacetabular blade of the ilium are identical to the much larger type of“Lythrosuchus” langstoni, suggesting that “Lythrosuchus” is a junior synonym ofPoposaurus. However, we retain the speciesPoposaurus langstoni nov. comb. on the basis of larger size, absence of a distinct ridge on the lateral surface of the ilium, and the lack of a pit on the proximal end of the ischium. A phylogenetic analysis suggests a monophyletic Poposauroidea and a paraphyletic “Rauisuchia”.     

Materials towards the revision of the genus Triplophysa rendahl, 1933 (Cobitoidea: Balitoridae: Nemacheilinae): A revision of nominal taxa of Herzenstein (1888) described within the species “Nemachilus” stoliczkae and “N.” dorsonotatus, with the description of the new species T. scapanognatha sp. nova

This study reviews the nominal taxa within the species “Nemachilus” stoliczkae and “N.” dorsonotatus, described by Herzenstein (1888). N. dorsonotatus, N. d. retropinnis, and N. plagiognathus are reviewed within the species Triplophysa stoliczkae; it is suggested that this species is represented by at least 5 subspecies. N. stoliczkae brevicauda, N. s. leptosoma (synonym, N. s. productus), and N. s. crassicauda are considered as valid species, the last 2 are related to the group including the species T. tenuis, T. choprai, T. tenuicauda, and T. yasinensis rather than with T. stoliczkae. It was shown that the “stoliczkae” species group (sensu Prokofiev, 2001) should be divided into three species groups: “stoliczkae” (s. str.), “tenuis,” and “robusta”; their diagnoses and the species composition are presented. A new species from the “stoliczkae” group is described (T. scapanognatha sp. nova), related to T. tanggulaensis (Zhu, 1982). Original Russian Text ? A.M. Prokofiev, 2007, published in Voprosy Ikhtiologii, 2007, Vol. 47, No. 1, pp. 5–25.     

Non-consumptive factors affecting foraging patterns in Antarctic penguins: a review and synthesis

Recent research has clearly shown that the fear of predation, i.e. aversion to taking risks, among mesopredators or grazers, and not merely flight from an apex predator to avoid predation, is an important aspect of ecosystem structuring. In only a few, though well-documented cases, however, has this been considered in the marine environment. Herein, we review studies that have quantified behavioral responses of Adélie penguins Pygoscelis adeliae and emperor penguins Aptenodytes forsteri to the direct presence of predators, and question why the penguins avoid entering or exiting the water at night. We also show, through literature review and new analyses of Adélie penguin diving data, that Antarctic penguins are capable of successful prey capture in the dark (defined here as <3.4 lux). Finally, we summarize extensive data on seasonal migration relative to darkness and prey availability. On the basis of our findings, we propose that penguins’ avoidance of foraging at night is due to fear of predation, and not to an inability to operate effectively in darkness. We further propose that, at polar latitudes where darkness is more a seasonal than a year-round, daily feature, this “risk aversion” affects migratory movements in both species, consistent with the “trade-off” hypothesis seen in other marine vertebrates weighing foraging success against predation risk in their choice of foraging habitat. Such non-consumptive, behavioral aspects of species interactions have yet to be considered as important in Southern Ocean food webs, but may help to explain enigmatic movement patterns and choice of foraging grounds in these penguin species.     

ECTOPARASITE LOAD IN A LAUGHING DOVE WITH A DEFORMED MANDIBLE

At Dyer Island, South Africa, observations of predation of seabirds by Cape Fur Seals Arctocephalus pusillus pusillus were made during 2004 and 2006/07. It was estimated that seals killed about 7% of adult African Penguins Spheniscus demersus annually. This may have contributed to the penguin colony not increasing in spite of food becoming more available in its vicinity during an eastward shift of epipelagic fish resources off South Africa at the start of the twenty-first century. Adult penguins were most susceptible to mortality in the breeding season as they returned to feed chicks in the evening. Seals also killed substantial numbers (3–9%) of Cape Cormorant Phalacrocorax capensis fledglings as they left the island, mostly in the morning. The losses of Cape Cormorant fledglings accounted for about 95% of the overall mortality of seabirds attributable to seals. Mortality inflicted by seals on adults of four species of cormorant was negligible and no predation by seals on gulls and terns was observed.     

PARTITIONING OF NESTING SPACE AMONG SEABIRDS OF THE BENGUELA UPWELLING REGION

Duffy, D. C. & La Cock, G. D. 1985. Partitioning of nesting space among seabirds of the Benguela upwelling region. Ostrich 56:186-201.

An examination of nesting habitats used by the four main species of seabirds nesting on southern African islands (Jackass Penguin Spheniscus demersus, Cape Cormorant Phalacrocorax capensis, Bank Cormorant P. neglectus and Cape Gannet Morus capensis) revealed relatively minor differences and extensive overlaps between species, primarily in subcolony size, steepness of nesting substratum, and proximity to cliffs. A weak dominance hierarchy existed; gannets could displace penguins, and penguins could displace cormorants. This hierarchy appeared to have little effect on partitioning of nesting space. Species successfully defended occupied sites in most cases of interspecific conflict, suggesting that site tenure by one species could prevent nesting by another. The creation of additional nesting space on Namibian nesting platforms did not increase guano harvests, suggesting that nesting space had not previously limited the total nesting population of Cape Cormorants, the most abundant of the breeding species, in Namibia.

While local shortages of nesting space may occur, populations of the four principal species of nesting seabirds in the Benguela upwelling region do not seem to have been limited by the availability of nesting space on islands.     

Comparative fuel metabolism in Gentoo and King Penguins: adaptation to brief versus prolonged fasting

To compare fuel utilization in large birds adapted to brief or prolonged fasting, protein and lipid utilization were quantified in the Gentoo Penguin (Pygoscelis papua) and the King Penguin (Aptenodytes patagonica). The inshore feeder Gentoo Penguin fasts for only a few days in its colony, while King Penguin chicks starve for several months in the subantarctic winter and male King Penguins starve for 5–6 weeks at the beginning of their breeding cycle. After an initial decrease in both daily body mass loss and nitrogen excretion during the first days of food deprivation, these two parameters thereafter stabilized at low values. At that time, protein utilization accounted for 15% of total energy expenditure in Gentoo Penguins and only 6% in King Penguin chicks during winter, the remainder (85% and 94%, respectively) being provided by fat oxidation. Similar percentages in fuel metabolism as seen in chicks during winter were reached in fasting adult King Penguins and spring chicks. However, a seasonal adaptation occurs in fasting chicks because energy expenditure is lower during winter. As previously described in starved mammals, the effectiveness in protein sparing could be related to the initial adiposity of the birds: the larger the fat stores (about 9% and 30% in Gentoo Penguins and winter chicks of King Penguins, respectively), the longer the fast duration and the better the level of protein conservation.     

7. SHORT NOTES FROM MEMBERS

Cooper, J., Ross, G. J. B. &; Shaughnessy, P. D. 1978. Seasonal and spatial distribution of Rockhopper Penguins ashore in South Africa. Ostrich 49:40-44. There are 30 records of Rockhopper Penguins Eudyptes chrysocome ashore in South Africa. Both the southern subspecies E.c. chrysocome, and the northern subspecies E.c. moseleyi have been recorded. The northern subspecies has occurred more frequently. Most records are of moulting juveniles in January and February. Records of adult birds are more scattered throughout the year. Rockhopper Penguins in South Africa have been recorded only south of 30S. Birds of the northern subspecies probably originate from South Atlantic islands (Tristan da Cunha group and Gough). Birds of the southern subspecies are probably from the Prince Edward Islands.     

Comparison of foraging strategies of incubating king penguins Aptenodytes patagonicus from Macquarie and Heard islands

The foraging strategies of king penguins from Heard and Macquarie islands were compared using satellite telemetry, time-depth recorders and diet samples. Trip durations were 16.8±3.6 days and 14.8±4.1 days at Macquarie and Heard islands, respectively. At Macquarie Island, total distances travelled were 1281±203 km compared to 1425±516 km at Heard Island. The total time the penguins spent at sea was 393±66 h at Macquarie Island and 369±108 h at Heard Island. The penguins from Macquarie Island performed more deep dives than those from Heard Island. King penguins from Macquarie Island travelled 1.5±0.2 km h⁻¹ day⁻¹ compared to 1.3±0.1 km h⁻¹ day⁻¹. At Macquarie Island, 19% of dives were upto 70–90 m depth compared to 35% at Heard Island. The main dietary prey species were the fish Krefftychthis anderssoni and the squid Moroteuthis ingens in both groups. The differences in the at-sea distribution and the foraging behaviour of the two groups of penguins were possibly related to differences in oceanography and bathymetric conditions around the two islands. Dietary differences may be due to interannual variability in prey availability since the two colonies were studied during incubation but in different years.     

The evolution of “egalitarian” and “despotic” social systems among macaques

Recent studies of captive macaques have revealed considerable inter-species differences in dominance styles among females. In “egalitarian” species such as stumptail (Macaca arctoides) or tonkean macaques (M. tonkeana), social interactions are more symmetrical and less kin-biased than in “despotic” species such as Japanese (M. fuscata) or rhesus macaques (M. mulatta). Field observations of moor macaques (M. maurus), close relatives of tonkean macaques, suggest that tolerance during feeding characterizes their egalitarian dominance style in the natural habitat. Although it has been proposed that communal defense against other groups may be the main selective force in the evolution of egalitarian dominance style among females, few field data support this prediction. A game theory analysis showed that both an “egalitarian” strategy and a “despotic” strategy are possible evolutionarily stable strategies (ESS) under certain conditions. The difference in dominance styles might reflect the difference in ESS. This means that an egalitarian dominance style can emerge without strong between-group contest competition. A phylogenetic comparison among macaques suggests that despotic dominance styles very likely evolved from egalitarian dominance styles. In the future, primate socioecological studies should pay more attention to the evolutionary history of each species.     

Upper limits of temperature for growth inChlorella (Chlorophyceae)

The upper limit of temperature for growth is a species-specific character in the genusChlorella. The limits of 14Chlorella species range from 26–30°C (C. saccharophila) to 38–42°C (C. sorokiniana), withC. fusca var.vacuolata (34°C) andC. kessleri (34–36°C) assuming an intermediate position. Thus, there is no wide gap in the temperature limits between the normal (“low-temperature”) species ofChlorella and the “high-temperature” species,C. sorokiniana.