WI - Cormorant Research Group Scientific Literature - Cormorants Last updated on 03-08-2010

Scientific literature on
Phalacrocoracidae and non-Palearctic species

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HOLLAND B.R., SPENCER H.G., WORTHY T.H. & KENNEDY M. 2010. Identifying Cliques of Convergent Characters: Concerted Evolution in the Cormorants and Shags. Syst. Biol., 59(4): 433–445. [Abstract. — A phylogenetic tree comprising clades with high bootstrap values or other strong measures of statistical support is usually interpreted as providing a good estimate of the true phylogeny. Convergent evolution acting on groups of characters in concert, however, can lead to highly supported but erroneous phylogenies. Identifying such groups of phylogenetically misleading characters is obviously desirable. Here we present a procedure that uses an independent data source to identify sets of characters that have undergone concerted convergent evolution. We examine the problematic case of the cormorants and shags, for which trees constructed using osteological and molecular characters both have strong statistical support and yet are fundamentally incongruent. We find that the osteological characters can be separated into those that fit the phylogenetic history implied by the molecular data set and those that do not. Moreover, these latter nonfitting osteological characters are internally consistent and form groups of mutually compatible characters or “cliques,” which are significantly larger than cliques of shuffled characters.We suggest, therefore, that these cliques of characters are the result of similar selective pressures and are a signature of concerted convergence.]

Yorio P., Copello S., Kuba L., Gosztonyi A. & Quintana F. 2010. Diet of Imperial Cormorants Phalacrocorax atriceps Breeding at Central Patagonia, Argentina. Waterbirds, 33: 70–78. [Abstract. — Diets of breeding Imperial Cormorants Phalacrocorax atriceps were studied at two breeding colonies, Islas Blancas and Isla Arce, located approximately 30 km apart in an area subject to increasing fishing pressure off Central Patagonia, Argentina. The goal was to assess differences between locations and diet variation among stages of the breeding cycle. Pellet casts (403 and 358 pellets per colony, respectively) were collected from November 2002 to February 2003. Analyses of the pellets revealed that Imperial Cormorants at Islas Blancas and Isla Arce fed on at least 25 and 23 prey types, respectively. Fish showed the highest frequency of occurrence at both colonies (> 70%), followed by crustaceans and molluscs. Of the fish prey, Merluccius hubbsi (22–71%), Engranlis anchoita (16–51%) and Raneya brasiliensis (5–48%) showed the highest frequencies of occurrence, depending on the colony and breeding stage. At Islas Blancas, the consumption of fish and crustaceans was similar among breeding stages (incubation, young chicks and old chicks), while it was significantly different at Isla Arce. Overall contribution by frequency of occurrence showed that M. hubbsi was the most frequent prey at Islas Blancas (58%), and E. anchoita and Pleoticus muelleri were more frequent at Isla Arce (48 and 45%, respectively). Also, overall contribution by mass of the main fish prey indicated differences between colonies. Given the commercial value of the main prey species, cormorant feeding requirements and spatial ecological needs should be included as considerations in coastal fisheries management and future development.]






HUGHES J. KENNEDY M., JOHNSON K.P., PALMA R.P. & PAGE R.D.M. 2007. Multiple Cophylogenetic Analyses Reveal Frequent Cospeciation between Pelecaniform Birds and Pectinopygus Lice. Syst. Biol., 56(2): 232–251. [Abstract. Lice in the genus Pectinopygus parasitize a single order of birds (Pelecaniformes). To examine the degree of congruence between the phylogenies of 17 Pectinopygus species and their pelecaniform hosts, sequences from mitochondrial 12S rRNA, 16S rRNA, COI, and nuclear wingless and EF1-a genes (2290 nucleotides) and frommitochondrial 12S rRNA, COI, and ATPases 8 and 6 genes (1755 nucleotides) were obtained for the lice and the birds, respectively. Louse data partitions were analyzed for evidence of incongruence and evidence of long-branch attraction prior to cophylogenetic analyses. Host-parasite coevolution was studied by different methods: TreeFitter, TreeMap, ParaFit, likelihood-ratio test, data-based parsimony method, and correlation of coalescence times. All methods agree that there has been extensive cospeciation in this host-parasite system, but the results are sensitive to the selection of different phylogenetic hypotheses and analytical methods for evaluating cospeciation. Perfect congruence between phylogenies is not found in this association, probably as a result of occasional host switching by the lice. Errors due to phylogenetic reconstruction methods, incorrect or incomplete taxon sampling, or to different loci undergoing different evolutionary histories cannot be rejected, thus emphasizing the need for improved cophylogenetic methodologies.]

Quintana F., Wilson R.P. & Yorio P. 2007. Dive depth and plumage air in wettable birds: the extraordinary case of the imperial cormorant. MEPS, 334: 299-310. [Abstract. Cormorants are considered to be remarkably efficient divers and hunters. In part, this is due to their wettable plumage with little associated air, which allows them to dive with fewer energetic costs associated with buoyancy from air in the feathers. The literature attributes particularly exceptional diving capabilities to cormorants of the ‘blue-eyed’ taxon. We studied the diving behaviour of 14 male imperial cormorants Phalacrocorax atriceps (included in the blue-eyed taxon) in Patagonia, Argentina, and found that this species did indeed dive deeper, and for longer, than most other non-blue-eyed cormorant species. This species also exhibited longer dive durations for any depth as well as longer recovery periods at the surface for particular dive durations. We propose that this, coupled with atypically long foraging durations at sea in cold water, suggests that cormorants of the blue-eyed complex have a plumage with a substantial layer of insulating air. This is given credence by a simple model. High volumes of plumage air lead to unusually high power requirements during foraging in shallow, warmer waters, which are conditions that tend to favour wettable plumage. However, deep dives and/or cold water should favour the blue-eyed phenotype, which explains their essentially high latitude distribution.]




Kalmbach E. & Becker P. H. 2005. Growth and survival of neotropic cormorant (Phalacrocorax brasilianus) chicks in relation to hatching order and brood size. Journal of Ornithology, 146(2): 91-98.



Gil de Weir K., Weir E., Casler C. & Aniyar S. Ecological Functions and Economic Value of the Neotropic Cormorant Phalacrocorax brasilianus in Los Olivitos Estuary, Venezuela. [Abstract. We determined the ecological function and economic value of a colony of piscivorous Neotropic Cormorants at the Los Olivitos Wildlife Refuge and Fisheries Reserve (WRFR), Lake Maracaibo, Venezuela. Colony size increased from 17,000 to approximately 40,000 in two years. Lake Maracaibo supports one of the most productive artesinal fisheries in Venezuela and the cormorant colony comprises 2 km of coastal mangrove. Neotropic Cormorants and fishermen use the same area, but do they compete? What is the ecological role of the Neotropic Cormorant in that area? To study the economic value of Neotropic Cormorants, we established ecological functions in the area of interest. Where, how many and how much do they feed? An ecological study of abundance, distribution and diet of Neotropic Cormorants was undertaken from 1999 to 2001. Abundance and distribution was discerned from monthly censuses and dietary composition was obtained via stomach and pellet analysis. An economic study was developed to estimate the economic impact and value of the Neotropic Cormorant population using four ecological-economic functions 1) Harvesting cormorants for food M(N), 2) Cormorants as contributors to fish diversity FD(N), 3) Cormorants as indicators of presence of fish schools S(N) and 4) Cormorants as contributors to fish biomass due to guano production B(GN). These functions were established after literature review and selection of goods, services, and attributes provided by Neotropic Cormorants in Los Olivitos Estuary and feasible for the study. The economic Total Value of the Neotropic Cormorant Population TV (N) was defined as the value of Cormorants to fishermen; changes in cormorant numbers would imply changes in the fishermen’s well-being. Ecological results indicated the population is increasing exponentially. Eighty-three percent of the population fed outside of the WRFR. Diet consisted mostly of 19 fish species in four families (Ariidae, Engraulidae, Gerreidae and Bothidae), and one shrimp. Monthly changes in dietary composition were observed. Average daily consumption was 225g, but before migration, birds may consume 800g/day. Based on the list of 9 commercial species consumed and fish size, no competition occurred. Estimated values of S(N), M(N) and FD(N) were positive, but B(GN) was negative.The Net Value of the Neotropic Cormorant population obtained only from S(N) + B(G,N) was $6,793,871/year. The Neotropical Cormorant population does not presently compete with artisanal fisheries in Lake Maracaibo, but if habitat is not a limiting factor and numbers of birds continue to increase, future conflicts could arise. This relationship offers unique opportunities to develop an integral ecological and economic dynamic model to study different scenarios to establish management policies for Neotropic Cormorants and artesinal fisheries, aquaculture, fish diversity and conservation.]


Gonçalves de Oliveira T. C. & Milléo Costa L. C. 2002. Space distribution of the Phalacrocorax brasilianus for occupation levels on the branches of the trees of the São Lourenço Park, Curitiba, Paraná, Brazil. Atualidades Ornitológicas (AO), n. 108, p.03, July/August. 2002.


Casaux R., Favero M., Silva P. & Baroni A. 2001. Sex differences in diving depths and diet of Antarctic Shags at the South Shetland Islands JOURNAL OF FIELD ORNITHOLOGY, 72(1): 22-29

Burger J. & Gochfeld M. 2001. Metal levels in feathers of cormorants, flamingos, and gulls from the coast of Namibia in southern Africa. Environmental Monitoring and Assessment, 69: 195-203.

Dorfman E. J. & Kingsford M. J. 2001. Environmental determinants of distribution and foraging behaviour of cormorants (Phalacrocorax spp.) in temperate estuarine habitats. Marine Biology, 138(1): 1-10. --- The distribution and behaviour of cormorants in esturarine environments was examined on the central coast of New South Wales, Australia, with respect to habitat associations at different spatial scales. No consistent variation in abundance was found for four species of cormorants (great Phalacrocorax carbo, pied P. varius, little back P. sulcirostris, and little pied P. melanoleucos) with state of tide (high and low) and time of day (early, middle, and late) at five stuarine locations. ---

Filardi C.E. & Rohwer S. 2001. Life history implications of complete and incomplete primary molts in Pelagic cormorants. The Condor, 103(3): 555-569. --- We describe the rules of primary flight-feather replacement for Pelagic Cormorants (Phalacrocorax pelagicus), and contrast the completeness of primary replacement in individuals from Asia and North America. In adult Pelagic Cormorants primary replacement is stepwise, with multiple waves of molt, each initiated at the innermost primary (P1), proceeding simultaneously toward the tip of the wing. Shugart and Rohwer's (1996) ontogenetic model for generating and maintaining stepwise primary replacement depended upon incomplete molts. In each new episode of molt, waves of primary replacement were thought to be initiated at P1 and at each arrested wave that had failed to replace all old feathers in the preceding molt. Because most adult Pelagic Cormorants from North America completely replace their primaries but maintain stepwise primary molts, the latter assumption must be relaxed. In contrast to the present-day situation in North America, Pelagic Cormorants from northeastern Asia have incomplete molts of their primaries, and may be forced to skip breeding in some years to clear their wings of overworn primaries. Young birds from Asia start the replacement of their juvenile primaries later than North American birds and replace more feathers simultaneously.

Frere, E. & Gandini P.A. 2001. Aspects of the breeding biology of the red-legged Cormorant Phalacrocorax gaimardi on the Atlantic coast of South America. Mar. Orn. 29: 67 - 70. [Abstract. Red-legged Cormorants Phalacrocorax gaimardi breed in Argentina, Chile and peru. In Argentina their breeding range is restricted to a short section of coastline in southern Patagonia. We studied two colonies located on high rocky cliffs, 2 to 4 m above the high tide line. At one colony, nests were protected from prevailing winds whereas at the other colony most of the nests were exposed. Of active nests, 15 % had two eggs, 66 % had three eggs, and 19 % had four eggs, to give a mean cluch size of 3.04 0.47. Egg dimensions were 60.3 2.4 x 37.1 1.4 mm. The incubation period ranged from 34 days to 38 days with chicks hatching from mid-November to the first week of December. Red-legged Cormorants lay more and smaller eggs than do those of two sympatric cormorant species, the Rock Cormorant P. magellanicus and the Imperial Cormorant P. atriceps, probably as a result of differences in foraging ranges. Avian predation on eggs seems to be an important mortality factor for this species and wind has also an important effect on breeding success, possibly exacerbating avian predation.]

Huner J. V. & Jeske C. 2001. Observations on the occurrence and food habits of double-crested cormorants and neotropic cormorants in south Louisiana crawfish ponds: The journal of Louisiana ornithology, 5(1): 22-30.

Kato A., Watanuki Y. & Naito Y. 2001. Foraging and breeding performance of Japanese cormorants in relation to prey type. Ecological Research, 16(4): 745-758. [Seabirds are high trophic predators in marine ecosystems and are sensitive to change in food supply and thus seabirds can be used as monitors of the marine environment. In order to study the foraging responses of Japanese cormorants Phalacrocorax filamentosus breeding at Teuri Island, Hokkaido to changes in fish availability, the diet was assessed from the regurgitations of parents and chicks, and diving behaviour was measured by using time-depth recorders. Breeding performance (brood size, chick growth, breeding success) was monitored using conventional methods to study their breeding responses. Japanese cormorants changed the diet and foraging behaviour over four summers. The birds fed mainly on epipelagic schooling fish when they were available and on demersal fish when pelagic fish availability was low. They tended to dive deeper and longer in a year when they fed mainly on demersal fish than the other years, reflecting the change in the depth distribution of prey fish. Chick growth rate did not differ among years, but fledging success was lower in the years of demersal fish as their meal delivery rate was low. When epipelagic schooling fish were considered scare, parents maintained chick growth by reducing brood size. High variability and unpredictability in pelagic fish abundance are key factors affecting the foraging and breeding performance of Japanese cormorants, which could potentially be used to monitor fish resources.]

Matthews D. 2001. SEABIRD SANCTUARY. Spurred by a cormorant with a broken wing, a young zoologist established a Florida hospital for battered birds that evolved into a leading wildlife rehabilitation organization. Wildlife Conservation, 104(5): 22-27.

Quintana F. 2001. Foraging behaviour and feeding locations of Rock Shags Phalacrocorax magellanicus from a colony in Patagonia, Argentina. Ibis, 143: 547-553.


Kato A., Watanuki Y., Nisiumi I., Kuroki M., Shaughnessy P. & Naito Y. 2000. Variation in foraging and parental behavior of King Cormorants. The Auk, 117: 718-730. [ We studied sexual and individual differences in foraging and parental behavior of King Cormorants (Phalacrocorax albiventer) rearing chicks were studied at subantarctic Macquarie Island. Females dived mainly in the morning and males in the afternoon. Five females were shallow divers (1.9 to 6.8 m) and seven were deep divers (19.6 to 28.0 m), but males dived even deeper (15.6 to 44.2 m) than both type of females. In King Cormorant, males were 16% heavier than females and the body mass was not different between deep divers and shallow divers of females. The percentage of bottom time to dive cycle was larger for shallow diving females (40 ± 13%) than males (26 ± 4%) and deep-diving females (27 ± 3%). Total daily dive time and bottom time did not differ between males, deep- and shallow-diving females, because shallow-diving females dived more frequently (211 ± 81 dives per day) than males (68 ± 21 dives per day) and deep-diving females (70 ± 7 dives per day). Meal delivery rate, trip duration and percentage of time at sea did not differ between males, deep- and shallow-diving females. Females, especially shallow divers, compensated their shallow and short dives with more frequent dives. Consequently male and female cormorants provisioned their chicks at a similar rate, in spite of large sexual and individual variation in foraging behavior.]

Kennedy M., R. D. Gray & Spencer H. G. 2000. The phylogenetic relationships of the shags and cormorants: can sequence data resolve a disagreement between behavior and morphology ? Molecular Phylogenetics and Evolution, 17(3): 345–359. (Available on-line at http://www.idealibrary.com). [Ref. 2000-8]

Powlesland R.G. & Luke I.J. 2000. Breeding biology of Little Shags (P. melanoleucos) at Lindale, Wellington. Notornis 47: 1-5.

Sievert R., Filardi C.E., Bostwick K.S. & Peterson T.A. 2000. A critical evaluation of Kenyon's shag (Phalacrocorax [stictocarbo] kenyoni). The Auk, 117(2): 308-320. --- We examine the validity of Phalacrocorax [Stictocarbo] kenyoni, which was described by Siegel-Causey (1991) from the Aleutian Islands using midden remains and existing skeletal specimens. We emphasize a morphometric evaluation of the taxon using 224 skeletal specimens of North Pacific cormorants, but we also evaluate the qualitative characters originally used to characterize P. kenyoni. Principal components and discriminant function analyses of 14 skeletal characters failed to support the validity of the species. Similarly, all seven of the character states that Seigel-Causey described as unique to P. kenyoni also were found in P. pelagicus and P. urile. Thus, the three type specimens of P. kenyoni appear to be P. pelagicus. Although we could not confirm the validity of P. kenyoni, our morphometric analyses revealed that P. pelagicus individuals from the central Aleutians are smaller than those from surrounding populations. Received 21 December 1998, accepted 20 July 1999.

Torres P.V., J. Schlatter, R. Montefusco, A. Revenga, J. Marin, F. Lamilla & Ramallo J. 2000. Infection by Contracaecum rudolphii (Nematoda : Anisakidae) in the Neotropic cormorant Phalacrocorax brasilianus, and fishes from the estuary of the Valdivia river, Chile. STUDIES ON NEOTROPICAL FAUNA AND ENVIRONMENT, 2000 - 35(2): 101-108 [Ref. 2000-6]


 Carney K. M. & Sydeman W. J. 1999. A review of human disturbance effects on nesting colonial waterbirds. Waterbirds, 22(1): 68-79.

Crawford R. J. M., Dyer B. M., Cordes I. & Williams A. J. 1999. Seasonal pattern of breeding, population trend and conservation status of bank cormorants Phalacrocorax neglectus off south western Africa. Biological Conservation, 87(1): pp. 49.

Dinsmore S. J. 1999. Neotropic Cormorant at Sutherland Reservoir. The Nebraska bird review, 67(2): 72.

Gerasimov N.N. 1999. [Pelagic Cormorant Phalacrocorax Cormorant Phalacrocorax pelagicus on the Karaginsky Island]. // The biology and conservation of the birds of Kamchatka. Moscow, 1: 83-86. In Russian with English summary. [rif. 1999-8]

Kato A., Watanuki Y., Shaughnessy P. D., Le Maho Y. & Naito Y. 1999. Intersexual differences in the diving behavior of foraging subantarctic cormorant (Phalacrocorax albiventer) and Japanese cormorant (P.filamentosus). C. R. Acad. Sci. Paris, Sciences de la vie/ Life Sciences, 322, 557-562. [Cormorants feed by feet-propelled diving. How cormorants optimize foraging is of a particular interest in relation to the understanding of the feeding strategies of diving birds, as well as within the debate about cormorants impact on sustainable resources. Using microdata loggers that recorded diving depth, we investigated the foraging strategy of males and females of subantarctic cormorants, which inhabit cold regions, and of Japanese cormorants, which live in the northern temperature zone. For both species, males and females daily spent the same amount of time submerged, and apparently captured the same amount of fish. However, males dived deeper and longer, which could be explained by their 15-20% larger body mass and may minimize potential competition for food.]

Lethaby N. & Moores N. 1999. Identification of Temminck's Cormorant. Dutch birding, 21(1): 1.

Nascetti G., Mattiucci S., Cianchi R., Berland B., Bullini & Paggi L. 2000. Genetic relationships among Contracaecum spp. (Nematoda: Anisakidae), parasites of cormorants and pelecans of the Boreal Region. In: 8th European Multicolloqium of Parasitology (EMOP VIII). Poznan, Polen, 10-14/9/2000. Acta Parasitologica 2000; 45(3):153.

Quintana F. 1999. Diving behavior of Rock Shags at a Patagonian colony of Argentina. Waterbirds, 22(3): 466-471.

Tovar S., Humberto. 1999. Desarollo poblacional de aves guaneras. Bol. Lima, no. 115: 101-112. --- The population development of the so-called guano birds (Guanay Cormorant Phalacrocorax bougainvillii, Peruvian Booby Sula variegata, Peruvian Pelican Pelecanus thagus) is analyzed for the period 1961 - 1989. In 1963 the bird population reached a maximum of 18 million. In 1966 descended to 4.7 million and stayed bellow 4 million in the following years. This was mainly caused by the industrial fish catch compounded in some years by the El Niño phenomena. Guano production was reduced accordingly from 250 metric tons in 1964 to 18.7 in 1989. ---

Zavalaga C.B. & Paredes R. 1999. Foraging behaviour and diet of the guanay cormorant. S. Afr. J. Mar. Sci., 21: 251-258.


Coldren M. K. et al. 1998. First Neotropic Cormorant, Phalacrocorax brasilianus (Aves: Phalacrocoracidae), breeding record for Arkansas. Southwest. Nat. 43: 496--498. (301 Krenek Tap #65, College Station, TX 77840, USA.) ---Breeding in mixed species heronry in Lafayette County in 1996.---

Hebschi, A. 1998. Foraging site preferences of Brandt's Cormorants off the Santa Cruz, California, coast. Colon. Waterbirds 21: 245--250. (Dept. Biol., Univ. Calif., Santa Cruz, CA 95064, USA; EM: ahebsch@prbo.org)---Phalacrocorax penicillatus foraging alone or in groups prefer areas less affected by heavy winds and seas; solitary foragers occur more often in rocky reef habitats than in habitats with sandy bottoms.---

Kato A., Watanuki Y. & Naito Y. 1998. Benthic and pelagic foraging of two Japanese cormorants, determined by similtaneous recording of location and diving activity. J. Yamashima Inst. Ornithol., 30: 101-108. [One male and one female Japanese cormorants, Phalacrocorax filamentosus, breeding at Terui Island, Hokkaido, foraged within 5 km of the island and in the strait between the island and mainland (17-27 km from the island). The foraging range of Japanese cormorants appeared to be limited to the neritic water between 10 and 60 m sea depth, where the seabed sediment was mainly rock and sand. Both birds showed high foraging site fidelity, with the male foraging to the south-east and the female to the north-east of the island. The male sometimes foraged in shallow water (<40 m), possibility feeding demersal fish. In deep water (>40 m), the male may have fed on epipelagic schooling fish. In contrast, the female did not reach the sea-bottom in deep or shallow waters, and it may have fed on epipelagic schooling fish.]

Hobson K.A. Pelagic Cormorant. (Phalacrocorax pelagicus). In: A. Poole & F. Gill (eds.), The Birds of North America Inc. No. 282, pp. 28.

Gawlik D. E. et al. 1998. Long-term trends in population and community measures of colonial-nesting waterbirds in Galveston Bay Estuary [Texas]. Colon. Waterbirds 21: 143--151. (Everglades Systems Res. Div., S. Florida Water Distr., 3301 Gun Club Rd., West Palm Beach, FL 33406, USA; EM: dale.gawlik@sfwmd.gov). --- From 1973-1990 populations of Roseate Spoonbill, Ajaia ajaja, Snowy Egret, Egretta thula, and Black Skimmer, Rynchops niger, decreased significantly; Neotropical Cormorants, Phalacrocorax brasilianus, and Sandwich Tern, Sterna sandvicensis increased. Changes thought due to loss of coastal marsh and increased feeding conditions outside Galveston Bay.---

Grémillet D. & Wilson R. P. 1998. A remote-controlled net trap for ground-nesting cormorants. Seabird, 20: 44-47.

Mills K.L. 1998. Multispecies seabird feeding flocks in the Galapagos Islands. The Condor, 100(2): 277-285. --- I examined the species composition, frequency, distance offshore, and duration of multispecies seabird feeding flocks in the Galapagos Islands, Ecuador. Flocks were comprised of Galapagos Penguins (Spheniscus mendiculus), Flightless Cormorants (Campsohaelius ANannopterumB harrisi), Brown Pelicans (Pelecanus occidentalis), Brown Noddies (Anous stolidus), Blue-footed (Sula nebouxii) and Masked Boobies (Sula dactylatra), Magnificent Frigatebirds (Fregata magnificens), and Audubon Shearwaters (Puffinus Iherminieri).

Quintana F. & Yorio P. 1998. Kelp Gull Larus dominicanus predation on an Imperial Cormorant Phalacrocorax atriceps colony in Patagonia. Mar. Orn., 26: 84-85.

Wallace E.A.H. & Wallace G.E. Brandt's Cormorant (Phalacrocorax penicillatus). In: A. Poole & F. Gill (eds.), The Birds of North America Inc., No. 362, pp. 28.


Albrieu C. & Navarro J. L . 1997. [Location and population size of cormorant colonies in the Deseado estuary, Santa Cruz, Argentina.] El Hornero 14(4): 243--246. ---Colonial nesting of Phalacrocorax gaimardi, Phalacrocorax magellanicus and Phalacrocorax olivaceus, with data on distribution and characteristics of the colonies and conservation.---

Dinsmore S. J. 1997. First record of a Neotropic Cormorant for Iowa. Iowa Bird Life, 67: 131-132. (6122 W. Magnolia, Fort Collins, CO 80521, USA). [a Phalacrocorax brasilianus photographed on 4 May 1996 in Decatur Co.]

Doherty J.L. & Brager S. 1997. The breeding population of Spotted Shags (Stictocarbo punctatus punctatus) on Banks Peninsula (New Zealand). Notornis, 44: 49-56.

Green K. 1997. Biology of the Heard Island Shag Phalacrocorax nivalis. 1. Breeding behaviour. Emu, 97: 60-66.

Green K. 1997. Biology of the Heard Island Shag Phalacrocorax nivalis. 2. Breeding. Emu, 97: 67-75.

Green K. & Williams R. 1997. Biology of the Heard Island Shag Phalacrocorax nivalis. 3. Foraging, diet and diving behaviour. Emu, 97: 76-83.

Grémillet D. 1997. Catch per unit effort, foraging efficiency, and parental investment in breedíng Great Cormorants (Phalacrocorax carbo carbo). ICES J. Mar. Sci., 54: 635-644.

Hobson K.A. Pelagic Cormorant. (Phalacrocorax pelagicus). In: A. Poole & F. Gill (eds.), The Birds of North America Inc. No. 282, pp. 28.

Jackson C. E. 1997. Fishing with cormorants. Arch. Nat. Hist., 24: 189-211.

Pence D. B. et al. 1997. New records of subcutaneous mites (Acari: Hypoderatidae) in birds, with examples of potential host colonization events. J. Med. Entomol. 34: 411--416. (Dept. Pathol., Texas Tech Univ., Health Sci. Ctr., Lubbock, TX 79430, USA.). --- Host records from 14 avian species, including herons, ibis, egrets, storks, pelicans, cormorants and puffins.---

Pence D. B & Newman S. 1997. Neottialges neopelagicus new species (Acari, Hypoderatidae) from the Pelagic cormorant (Aves, Phalacrocoracidae, Pelecaniformes). Journal of Medical Entomology, 34(1): 33-37.

Siegel-Causey D. 1997. Molecular variation and biogeography of Rock Shags. Condor, 99: 139-150.


Barera-Oro E.R. & Casaux R.J. 1996. Fish as a diet of the Blue-eyed Shag at Halfmoon Island, South Shetland Islands. Cybium, 20: 37-45.

Dorfam E.J. & Read J. 1996. Nest predation by corvids on cormorants in Australia. Emu, 96: 132-135.

Kato A., Y. Naito, Y. Watanuki & Shaughnessy P. D. 1996. Diving pattern and stomach temperatures of foraging King cormorants at sub-Antarctic Macquarie Island. Condor, 98(4): 844-848.

Kopij G. 1996. Breeding and feeding ecology of the Reed Cormorant Phalacrocorax africanus in the Orange Free State, South Africa. Acta Ornithol. (Warsaw) 31: 89--99. (Dept. Zool. & Entomol., Univ. Orange Free State, P.O. Box 339, Bloemfontein 9300, South Africa). ---Information on hatching and fledging success, adult activity patterns and diet.---

Lea S. E. G. et al. 1996. Diving patterns in shags and cormorants (Phalacrocorax): tests of an optimal breathing model. Ibis, 138: 391--398. (Dept. Psychol., Univ. Exeter, Washington Singer Lab., Exeter EX4 4QG, UK.) --- Comparative study between Phalacrocorax aristotelis, Phalacrocorax melanoleucos, Phalacrocorax carbo and Phalacrocorax varius.---

Malacalza V.E. & Navas J.R . 1996. Biologia y ecologia reproductiva de Phalacrocorax albiventer (Aves: Phalacrocoracidae) en Punta Leon, Chubut, Argentina. Orn. Neotrop, 7: 53-61.

Reese P.R., Powlesland R. & Cornick D. 1996. Nesting of the Pied Shag (Phalacrocorax varius) at Makara Beach, Welllington (New Zealand). Notornis, 43: 212-213.

Spencer H.G., M. Kennedy & Gray R. D. 1996. An observation of aggressive nest defence in the Pitt Island Shag (Stictocarbo featherstoni). Notornis, 43: 208-210.

Taylor M. J. 1996. A Pied Shag (Phalacrocorax varius) beating the surface of the water while fishing. Notornis, 43: 211.

Watanuki Y. Kato A. & Naito Y. 1996. Diving performance of male and female Japanese cormorants. Canadian Journal of Zoology, 74: 1098-1109. [Sexual differences in the diving behavior of the sexually dimorphic Japanese Cormorant, Phalacrocorax capillatus (males are 26% heavier than females), were studied at Terui Island, Hokkaido, using time-depth recorders. A typical dive cycle involved a rapid descent phase, a bottom phase where they remained for a while, an ascent phase, and postdive surface phase. Depth and duration across individual birds were greater for males (15.1 ± 3.7(mean ± SD) m,37 ± 5 s, respectively) than those for females (7.2 ± 2.4 m , 24 ± 4 s, respectively). While submerged, females spent a similar propotion of time during the bottom phase to males, hence foraging efficiency (propotion of time at the bottom to total dive cycle time) did not differ berween the sexes. No sexual differences were found in descent and ascent rates, dive bout duration, or time spent underwater per day. No significant effects of dive duration on postdive surface time were observed for either sex, indicating that birds dived within an aerobic dive limit. However, mean dive durations and maximum dive durations for individual birds were a function of body mass to the power 1.49 and 1.87, respectively, suggesting that body mass partly constrains the diving behavior of this opportunistically feeding cormorant.]

Wanink J. H. 1996. Foraging locations of kingfishers and cormorants at Lake Victoria depend on the distribution of harvestable prey. Afr. J. Ecol., 34: 90-93.

Wilson R. P. & Grémillet D. 1996. Body temperatures of free living African penguins (Spheniscus demersus) and Bank cormorants (Phalacrocorax neglectus). Journal Experimental Biology, 199(10): 2215-2223.


Emslie S.D. 1995. A Catastrophic Death Assemblage of a New Species of Cormorant and Other Seabirds from the Late Pliocene of Florida. Journal of vertebrate paleontology, 15(2): 313-

Gandini P. & Frere E. 1995. Distribucion, abundancia, y ciclo reproductivo del cormoran gris Phalacrocorax gaimardi en la costa Patagonica, Argentina. Hornero, 14: 57-60.

Gosselin, M. 1995. [A poorly known bird family: Phalacrocoracidae.] Québec Oiseaux 6(2): 18--20. (Can. Mus. Nat., P.O. Box 3443, Station D, Ottawa, ON K1P 6P4, Can.)---General presentation of N. American cormorants. (French.)---

Grémillet D. D. 1995. "Wing-drying" in cormorants. J. Avian Biol., 26: 176.

Hustler K. 1995. Cormorant and Darter prey size selection under experimental conditions. Ostrich 66: 109--113. (P.O. Box 159, Victoria Falls, Zimbabwe).

Kasoma P. M. B. 1995. Cormorant regurgitation. and scavenging by large water birds at an avian loafing site in Queen Elizabeth National Park, Uganda. Afr. J. Ecol., 33: 294-296.

Keith J. A. 1995. Management policies for cormorants in Canada. Colonial Waterbirds, 18: 234-237.

Monadjem A. & Owen-Smithy R.N. 1995. The influence of water depth on foraging location of two species of kingfisher and two species of cormorant on the Sabie River within the Kruger National Park. Afr. J. Ecol. 33: 81-83.

Telfair II R. C. 1995. Neotropic Cormorant (Phalacrocorax brasilianus) population trends and dynamics in Texas. Bull. Texas Ornithol. Soc. 28: 7--16. (Texas Parks & Wildl. Dept., Manage. & Res. Stn., 11942 FM 848, Tyler, TX 75707, USA.)---After drastic population declines in the 1960's, populations have steadily increased. Banding data yield additional information.---

Telfair II R. C. & Morrison M. L 1995. Neotropic Cormorant Phalacrocorax brasilianus. Pp. 1-24 in: The birds of North America, No. 137. Poole A. & Gill F. (eds.), Academy of Natural Sciences, Philadelphia, Pennsylvania, and AOU, Washington, D.C., USA.

Valle C. A. 1995. Effective population size and demography of the rare Flightless Galapagos cormorant. Ecological Applications, 5(3): 601-617.

Wilson R. P. & Wilson M. P . 1995. Buoyancy and depth utilisation in foraging Cormorants: wet feathers and that sinking feeling. Gerfaut, 85: 41--47. (Inst. Meereskunde, Dusternbrooker Weg 20, D-24105, Germany.)---Phalacrocorax carbo, Phalacrocorax neglectus, Phalacrocorax capensis, Phalacrocorax coronatus.---


Malacalza V.E., T.I. Poretti & Bertelloti N.M. 1994. La dieta de Phalacrocorax albiventer en Punta Leon (Chubut, Argentina) durante la temporada reproductiva. Orn. Neotrop, 5: 91--97.

Sheppard Y. 1994/5. Cormorants and pelicans. Birds of the Wild 3(4):30-35.

van Tets G. F. 1994. Do cormorants eat freshwater mussels ? If not what does ? Emu, 94: 127-128.


Casaux R. J. & Barreraoro E. R. 1993. The diet of the Blue-eyed shag, Phalacrocorax atriceps bransfieldensis feeding in the Bransfield Strait. Antarctic Science, 5(4): 335-338.

Lefeure C. 1993. Las aves en los yacimientos del archipiélago del Cabo de Hornos y del Seno Grandi. [Birds from the graves of Cabo de Hornos and Seno Grandi.] Ans. Inst. Patagonia Ser. Cs. Hs., 22: 123-136. (URA 1415 du CNRS, Mus. Natl. d'Hist. Nat., 55 rue Buffon, 75005 Paris, France.)---Most abundant fossils at these sites on the southern coast of Chile are albatrosses, shearwaters, cormorants, and their allies.---

Martin P. (ED.). 1994. WHITEBREASTED CORMORANTS & KELP GULLS. Bee-eater; 45(4): 57.

Spear L. B. 1993. Dynamics and effect of Western gulls feeding in a colony of Guillemots and Brandts Cormorants. Journal of Animal Ecology, 62(3): 399-414.

Symens P., van den Berg A.B. & Keijl G.O. 1993. Morphology and distribution of Socotra Cormorant in Saudi Arabia. Dutch birding, 15(5): 215-


Bamber T. B. 1992. Flight speed of cormorant. Br. Birds 85: 237.

Chapdelaine G. & Brousseau P. 1992. Distribution, abundance and changes of seabird populations of the Gaspé Peninsula, Quebec, 1979 to 1989. Can. Field-Nat., 106: 427-434.

Douthwaite R. J. et al. 1992. DDT residues and mercury levels in Reed Cormorants on Lake Kariba, Zimbabwe: Hazard Assessment. Ostrich, 63: 123-127. (Tsetse Trypanosomiasis Control Br., P.O. Box 8283, Causeway, Zimbabwe.).

Humphries P., G. A. Hyndes and Potter I. C. 1992. Comparisons between the diets of distant taxa (Teleost and Cormorant) in an Australian estuary. Estuaries, 15: 327-334.

Hustler K. 1992. Buoyancy and its constraints on the underwater foraging behaviour of Reed Cormorants Phalacrocorax africanus and Darters Anhinga melanogaster. Ibis, 134: 229-236.

Kato, A., J.P. Croxall, Y. Watanuki and Y. Naito. 1992. Diving patterns and performance in male and female blue-eyed cormorants Phalacrocorax atriceps at South GeorgiaMarime Ornithology, 19: 117-129. [Diving pattern and performance in male and female Blue-eyed Cormorants Phalacrocorax atricepes were recorded with continuous-recording time-depth recorders for 10-13 days during the 1990 breeding season. For the female only data on time and depth were retrieved; diving was distributed through the day (including 31% at night); mean dive depth was 63 m. Male dived mainly in the afternoon and evening. Their mean dive depths were 61.4 and 83.9 m; mean dive durations 3.4 and 4.6 min; time at maximum depth (bottom time) averaged 28 and 44% of time submerged; descent time (0.4 and 0.5 min) was shorter than ascent time (1.6 and 2.9 min); dive/pause rations were 0.4 and 0.5 and surface interval was with preceding dive duration. The incubating male making one long dive bout per day, made fewer, longer dives, with less overall bottom time, than the chick-rearing male, which made at least two bouts per day. Most characteristics of diving in 1990 (a year of diving breeding success) were similar to data from 1989 (a year of breeding failure). However there were more, shorter dive bouts in 1989 and shallow travelling) dives were more frequent; both consistent with lower preyavalibility in 1989 necesstating frequent changes of forging sites.]

Livezey B.C. 1992. Flightlessness in the Galapagos cormorant (Compsohalieus [Nannopterum] harrisi): heterochrony, giantism and specialization. Zoological journal of the Linnean Society, 105(2): 155-

Melrose W.D. & Nicol S.C. 1992. Haematology, red cell metabolism and blood chemistry of the black-faced cormorant Leucocarbo fuscescens. Comparative biochemistry and physiology. A. 102(1): 67-

Wilson R. P., K. Hustler, P. G. Ryan, A. E. Burger & Noldéke E. C. 1992. Diving birds in cold water: do Archimedes and Boyle determine energetic costs? Am. Nat., 140: 179-200.


Ball D. 1991. A high incidence of bent beaks in nestling Pied Cormorants. Emu, 91: 257.

Croxall J P., Y. Naito, A. Kato, P. Rothery and Briggs D. R. 1991. Diving patterns and performance in the Antarctic Blue-Eyed Shag Phalacrocorax atriceps. J. Zool., 225: 177-199.

Siegel-Causey D. 1991. Systematics and biogeography of north Pacific shags, with a description of a new species. Occasional papers of the Museum of Natural History (Lawrence, Kansas: The University of Kansas), 140 : 1-17

Siegel-Causey D., C. Lefevre & A.B. Savinetskii. 1991. Historical diversity of cormorants and shags from Amchitka Island, Alaska. Condor, 93: 840-852.

Malacalza V. E. 1991. External characters in the offspring resulting from cross-breeding between cormorant species. Colonial Waterbirds, 14:180-83.

Rasmussen P. C. 1991. Relationships between South American king and blue-eyed shags. Condor, 93:825-39.

Siegel-Causey D. 1991. Systematics and biogeography of North Pacific shags, with a description of a new species. U. of Kans. Mus. Nat. Hist. Occ. Pap. 140:1-15.

Wanless S. and Harris M. P. 1991. Diving patterns of full-grown and juvenile Rock shags. Condor, 93: 44-48.


Barrett R. T., N. Rov and Montevecchi W. A. 1990. Diets of shags Phalacrocorax aristotelis and cormorants P. carbo in Norway and possible implications for ganoid stock recruitment. Marine Ecology Progress Series, 66: 205-218.

Boekelheide R. J., D. G. Ainley, H. R. Huber and Lewis T. J . 1990. Pelagic cormorant and double-crested cormorant. Pp. 195-217, in: D. C. Ainley and R. J. Boekelheide (eds.), Seabirds of the Farallon Islands: ecology, dynamics and structures of an upwelling-system community. Stanford Univ. Press, Palo Alto.

Green K., Williams R. & Slip D.J. 1990. Diet of the Macquarie Island cormorant Phalacrocorax atriceps purpurascens. Corella Serial 14(2): 53-55.

Green K., Williams R., Woehler E.J., Burton H.R., Gales N.J. & Jones R.T. 1990. Diet of the Heard Island cormorant Phalacrocorax atriceps nivalis. Antarctic Science Serial 2(2): 139-141.

Rosenberg D. K., C. A. Valle, M. C. Coulter & Harcourt S. A.. 1990. Monitoring Galapagos penguins and flightless cormorants in the Galapagos islands. Wilson Bull., 102(3): 525-32.

Sibley C. G. & Monroe B. L. Jr. 1990. Distribution and taxonomy of birds of the world. Yale Univ. Press, New Flaven.


Blaber, S. and T J. Wassenberg 1989. Feeding ecology of the piscivorous birds Phalacrocorax varius, P. melanoleucos and Sterna bergii in Moreton Bay, Australia: diets and dependence on trawler discards. Marine Biology, 101: 1-10.

Boekelheide R. J. and Ainley D. G . 1989. Age, resource availability and breeding effort in Brandt's cormorant. Auk, 106: 389-401.

Browning M. R. 1989. The correct name for the Olivaceous cormorant, "Maiague" of Piso (1658). Wilson Bull., 101(1): 101-106. --- William Pies' description in 1658 of the "Maiague" is shown to be a reference to the Olivaceous Cormorant. Phalacrocorax brasilianus should be reinstated as the correct name, with the type locality as eastern Brazil.

Cobley N. 1989. Aspects of the survival of blue-eyed shags (Phalacrocorax atriceps King). Antarctic Spec. Top., 1989: 93-96.

Rasmussen P. C. 1989. Post-landing displays of Chilean blue-eyed shags at a cliff-nesting colony. Bird Behav., 8: 51-54.

Trayler K. M., D. J. Brothers, R. D. Wooller & Potter I. C. 1989. Opportunistic foraging by three species of cormorants in an Australian estuary. J. Zool. Lond., 218: 87-98.


Dostine P. L. and Morton S. R . 1988. Notes on the food and feeding habits of cormorants on a tropical flood-plain. Emu 88:263-66.

Espitalier-Noel C., N. J. Adams and Mages N. T . 1988. Diet of the imperial cormorant Phalacrocorax atriceps at subAntarctic Marion Island. Emu 88:43-46.

Hobson K. A. & Carter H. R. 1988. Bill deformity in a Brand's cormorant chick. Calif. Fish Game, 74: 184-185.

Humphrey P. S., P. C. Rasmussen and Lopez N . 1988. Fish surface activity and pursuit-plunging by olivaceous cormorants. Wilson Bull. 100:327-28.

Malacaza V. E. and Hall M. A. 1988. Sexing adult King cormorants Phalacrocorax albiventer by discriminant analysis. Colonial Waterbirds, 11: 32-37.

Rasmussen P. C. 1988. Variation in the juvenal plumage of the red-legged shag (Phalacrocorax gaimardi) and notes on behavior of juveniles. Wilson Bull., 100:535-44.

Rasmussen P. C. and Hurnphrey P. S. 1988. Wing-spreading in Chilean blue-eyed shags (Phalacrocorax atriceps). Wilson Bull., 100:140-44.

Siegel-Causey D. 1988. Phylogeny of the Phalacrocoracidae. Condor, 90(4): 885-905.

Vermeer K., K. H. Morgan and Smith C. E. J. 1988. Population trends and nesting habitat of Double-crested and Pelagic cormorants in the Strait of Georgia. Pp. 94-98, in F. Vermeer and R. W. ButIer (eds.), The ecology and status of marine and shoreline birds in the Strait of Georgia, British Columbia. Can. Wildl. Serv., Ottawa.

Wilson R. P. & Wilson M. P. T. 1988. Foraging behaviour in four sympatric cormorants. Journal of Animal Ecology, 57(3): 943-955.


Anderson I. 1987. Epidemic of bird deformities sweeps U.S. New Scientist 3 Sept. 1987, 21.

Cooper J. 1987. Biology of the bank cormorant. 5. Clutch size, eggs and incubation. Ostrich 58:1-8.

Dowding J. E. and Taylor M. J . 1987. Genetics of polymorphism in the little shag. Notornis 34:51-57.

Drummond H. 1987. A review of parent-offspring conflict and brood reduction ìn the Pelecanìformes. Colonial Waterbirds, 10: 1-15.

Duffy C. D., R P. Wilson and Wilson M P. 1987. Spatial and temporal patterns of diet in the Cape Cormorant off southern Africa. Condor, 89: 830-834.

Pemberton D. and Gales R. P. 1987. Notes on the status and breeding of the imperial cormorant Phalacrocorax atriceps at Heard Island. Cormorant, 15:33-39.

Rasmussen P. C. 1987. Molts of the rock shag and new interpretations of the plumage sequence. Condor, 89:760-66.

Rijke A. M. 1987. The water repellency of water-bird feathers. Auk 104: 140-142.

Rosenberg D. K. and Harcourt S. A.. 1987. Population sizes and potential conservation problems of the endemic Galapagos penguin and flightless cormorant. Noticias de Galapagos, 45:24-25.

Schreiber R. W. and Clapp R. B. 1987. Pelecaniform feeding ecology. Pp. 173-88, in: J. P. Croxall (ed.), Seabirds. Feeding ecology and role in marine ecosystems. Cambridge Univ. Press, Cambridge.

Siegel-Causey D. 1987. Behaviour of the red-footed cormorant (Phalacrocorax gaimardi). Notornis, 34:1-9.

Taylor M. J. 1987. A colony of the little shag and the pied shag in which the plumage forms of the little shag freely interbreed. Notornis, 34:41-50.

Telfair R. C. II and Swepston D. A. 1987. Analysis of banding and marking nestling Anhingas, Olivaceus cormorants, Roseate spoonbills, Ibises, Bitterns, Herons and Egrets in Texas (1923-1983). Texas Parks and Wildlife Dep. Fed. Aid. Proj. W-103-R. Pwd-Bk-7100-152-7/87, Pp. 51.

Williams A. J. 1987. New seabird breeding localities, and an extension of bank cormorant range, along the Namib coast of southern Africa. Cormorant, 15:98-102.


Becker J. J. 1986. Reidentification of "Phalacrocorax" subVolans Brodkorb as the earliest record of Anhingidae. Auk 103:804-8.

Cooper J. 1986. Diving patterns of Cormorants Phalacrocoracidae. Ibis, 128: 562-570.

Cooper J. 1986. Biology of the bank cormorant. 4. Nest construction and characteristics. Ostrich 57:170-79.

Donnelly B. G. & Hustler K. 1986. Notes on the diet of the reed cormorant and the darter on the Lake Kariba during 1970 and 1971. Arnoldia (Zimbabwe), 9(24): 319-324.

Duffy D. C., R. P. Wilson, M. -P. Wilson and Velasquez C. R. 1986. Plunge-diving by olivaceous cormorants in Chile. Wilson Bull. 98:607-8.

Loutit R., D. Boyer and Brooke R. K . 1986. Cape cormorant (Phalacrocorax capensis) and jackass penguin (Spheniscus demersus) breeding colonies on the Narnibian mainland coast around Sylvia Hill. Cormorant, 13:185-87.

Mathews C. W. and Fordham R. A . 1986. Behaviour of the little pied cormorant (Phalacrocorax melanoleucos). Emu, 86:118-21.

Rasmussen P. C. 1986. Reevaluation of cheek patterns of juvenal-plurnaged blue-eyed and king shags. Condor, 88:393-95.

Shaw P. 1986. Factors affecting the breeding performance of Antarctic blue-eyed shags Phalacrocorax atriceps. Ornis Scand., 17:141-50.

Siegel-Causey D. 1986. Behavior and affinities of the Magellanic cormorant. Notornis, 33:249-57.

Siegel-Causey D. 1986. The courtship behavior and mixed-species pairing of king and imperial blue-eyed shags (Phalacrocorax albiventer and P. atriceps). Wilson Bull,. 98:571-80.

Siegel-Causey D. & Hunt C. L. Jr. 1986. Breeding-site selection and colony formation in double-crested and pelagic cormorants. Auk, 103:230-234.


Bernstein N. P. & Maxson S. J. 1985. Reproductive energetics of Blue-eyed Shags in Antarctica. Wilson Bull., 97: 450-462.

Brothers N. P. 1985. Breeding biology, diet and morphometrics of the king shag Phalacrocorax albiventer purpurascens at Macquarie Island. Aust. Wildl. Res. 12:81-94.

Cooper J. 1985. Foraging behaviour of non-breeding Imperial cormorants at the Prince Edward Islands. Ostrich, 56: 96-100.

Cooper J. 1985. Biology of the bank cormorant. 2. Morphometrics, plumage, bare parts and moult. Ostrich 56:79-85.

Cooper J. 1985. Biology of the bank cormorant. 3. Foraging behavior. Ostrich 56:86-95.

Cracraft J. 1985. Monophyly and phylogenetic relationships of the Pelecaniformes: a numerical cladistic analysis. Auk 102:834-53.

Hobson K. A. & Sealy S. G. 1985. Diving rhythms and diurnal roosting time of pelagic Cormorants. Wilson Bull., 97: 116-119.

Hogan G. G. 1985. Noosing adult cormorants for banding. N. Am. Bird Banding, 10(3): 76-77.

McNeil R., J.R. Rodriguez & Quellet H. 1985. Bird mortality at a power transmission line in northeastern Venezuela. Biological Conservation, 31: 153-165. [Casualties from distribution line collisions are considered significant. The authors suggest placing lines parallel rather than across flight paths; attempting to make lines more visible by luminous orange markings (wrapped tapes or clipped strips); and burying cables. Species recovered in this study included brown pelican, royal tern, black-crowned night heron, and neotropic cormorant. Data on flight patterns, behavior, feeding, and nesting are included. Frequency of casualties is said to be related to species, composition, behavior patterns, flight characteristics or flight directions, and local features.]

Nakagawa H. 1985. [Breeding of Temminck's cormorant Phalacrocorax filamentosis in the Shiretoko Peninsula, Hokkaido.] Bull. Shiretoko Mus., 7:71-72. (Japanese, English summary).

Ryan P. C. & Hunter S . 1985. Early breeding of imperial cormorants Phalacrocorax atriceps at Prince Edward Island. Cormorant, 13:31-34.

Shaw P. 1985. Age-differences within breeding pairs of blue-eyed shags Phalacrocorax atriceps. Ibis, 127:53743.

Shaw P. 1985. Brood reduction in the blue-eyed shag Phalacrocorax atriceps. Ibis, 127:476-94.


Barlow C. G. and Bock K. 1984. Predation of fish in fish dams by Cormorants, Phalacrocorax ssp. Austr. Wildl. Res., 11: 559-566.

Bernstein, N P. and S J. Maxson 1984. Sexually distinct daily activity patterns of Blue-eyed Shags in Antarctica. Condor, 86: 151-156.

Carter H. R., K. R. Hobson and Sealy S. G. 1984. Colony site selection by pelagic cormorants (Phalacrocorax pelagicus) in Barkely Sound, British Columbia. Colonial Waterbirds 7:25-34.

Elowson A. M. 1984. Spread-wing posture and the water-repellency of feathers: a test of Rijke's hypothesis. Auk, 101: 371-383.

Hennemann W. W. 1984. Spread-winged behaviour of Doubled-crested and Flightless cormorants Phalacrocorax auritus and P. harrisi: wing drying or thermoregulation? Ibis, 126: 230-239.

Mahoney S. A. 1984. Plumage wettability of aquatic birds. Auk, 101:181-85.

Malacalza V. E. 1984. Biologia reproductiva de Phalacrocorax albiventer. I. Nidificacion en Punta Tombo. Contr. 98, Cent. Nac. Pata., ConseJ. Nac. de Invest. Cient. y Tec.

Olver M. D. 1984. Breeding biology of the reed cormorant. Ostrich, 55:133-40.

Talent L. 1984. Food habits of wintering Brandt's cormorants. Wilson Bull., 96:130-134.

Tindle R. 1984. The evolution of breeding strategies in the flightless cormorant (Nannopterum harrisi) of the Galapagos. Biol. J. Linn. Soc., 21:157-164.

Vermeer K. and Rankin L. 1984. Population trends in nesting Double-crested and Pelagic cormorants in Canada. Murrelet, 65:1-9.


Duffy D. C. 1983. Competition for nesting space among Peruvian guano birds. Auk 100:680-88.

Duffy D. C. & Laurenson L. 1983. Pellets of Cape Cormorant as indicators of diet. Condor, 85: 305-307.

LIewellyn L. C. 1983. Movements of cormorants in south-eastern Australia and the influence of floods on breeding. Aus. Wíldl. Res., 10:149-67.

Matthews J. P. and Berruti A . 1983. Diet of Cape gannet and Cape cormorant off Walvis Bay, 1958-59. S. Afr. J. Mar. Sci., 1:61-63.

Trillmich F., K. Trillmich, D. Limberger and Arnold W. 1983. The breeding season of the flightless cormorant Nannopterum harrisi at Cabo Hammond, Fernandina. Ibis, 125:221-223.

Winkler H. 1983. The ecology of Cormorants (Genus Phalacrocorax) development. Hydrobiologia, 12: 192-199.

Williams A. J. and Cooper J. 1983. The Crowned cormorant: breeding biology, diet and offspring-reduction strategy. Ostrich 54:213-19.


Bernstein N. P. & Maxson S. J. 1982. Behaviour of the Antarctic blue-eyed shag Phalacrocorax atriceps bransfieldensis. Notornis 29:197-207.

Brooke R. K., Cooper J., P. A. Shelton & Crawford R. J. M. 1982. Taxonomy, distribution, population size, breeding and conservation of the Whitebreasted cormorant, Phalacrocorax carbo, on the Southern African coast. Gerfaut 72: 189-220.

Clapp R. B., M. K. Klimkiewicz & Kennard J. H. 1982. Longevity records of North American birds: Gaviidae through Alcidae. J. Field Ornithol., 53: 81124.

Cooper J., R. K. Brooke, P. A. Shelton and Crawford R. J. M. 1982. Distribution, population size and conservation of the Cape cormorant Phalacrocorax capensis. Fish. Bull. S. Afr. 16:121-43.

Crawford R. J. M., P. A. Shelton, R. K. Brooke & Cooper J . 1982. Taxonomy, distribution, population size and conservation of the Crowned cormorant Phalacrocorax coronatus. Gerfaut, 72: 3-30.

Foster M. S. & Fitzgeral L. A. 1982. A technique for live-trapping Cormorants. J. Field Orn., 53(4): 422-423.


Newell J. G. and Sutton C. M.. 1982. The olivaceous cormorant in Oklahoma. Bull. Okla. Ornith. Soc., 15:1-5.

Telfair R. C. II, C. D. Frentress & Davis B. G. 1982. Food of fledgling Olivaceus cormorants (Phalacrocorax olivaceus) in east central Texas. Bull. Texas Ornith. Soc., 15: 16-18.


Ainley, D., T. Anderson and P R. Kelley 1981. Feeding ecology of marine cormorants in Southwestern North America. Condor, 83: 120-131.

Cooper J. 1981. Biology of the bank cormorant. l. Distribution, population size, movements and conservation. Ostrich 52: 208-215.

Siegel-Causey D. and Hunt C. L. Jr. 1981. Colonial defense behavior in double-crested and pelagic cormorants. Auk, 98:522-531.


Miller B. 1980. Ecology of the little black cormorant Phalacrocorax sulcirostris and little pied cormorant, P. melanoleucos in inland New South Wales. Il. Proximate control of reproduction. Aust. Wildl. Res., 7:85-101.

Ono K. 1980. Comparative osteology of three species of Japanese cormorants of the genus Phalacrocorax (Aves, Pelecaniformes). Bull. Nat. Sci. Mus. Japan, Ser. C (Geol. & Paleontol.), 6:129-51.


1970 - 1979

Berry H. H. 1974. The crowned race of reed cormorant Phalacrocorax africanus coronatus breeding underneath Walvis Bay guano platform, South West Africa. Madoqua 8:59-62.

Berry H. H. 1974. The Crowned race of Reed cormorant Phalacrocorax africanus coronatus breeding underneath Walvis Bay guano platform, South West Africa. Madoqua ser. I, 8: 59-62.

Berry H. H. 1976. Physiological and behavioral. ecology of the Cape cormorant Phalacrocorax capensis. Madoqua, 9(4): 5-55.

Berry H. H. 1977. Seasonal fidelity of Cape cormorants to nesting areas. Cormorant, 2: 5-6.

Birkhead M. E. 1978. Some aspects of the feeding ecology of the Reed cormorant and Darter in Lake Kariba, Rhodesia. Ostrich, 49: 1-7.

Blankley W. O. 1981. Marine foods of kelp gulls, lesser sheathbills, and imperial cormorants at Marion Island (subantarctic). Cormorant 9:77-84.

Chapdelaine G. 1977. Le Grand Cormoran (Phalacrocorax carbo) en hiver, le long des cótes de la Péninsule de Gaspésie, Québec. Can. Field-Nat., 91: 184.

Conroy J. W. H. & Twelves E. L. 1972. Diving depths of the gentoo penguin (Pygoscelis papua) and the blue-eyed shag (Phalacrocorax atriceps) for the South Orkney Islands. Brit. Antarct. Survey Bull. 30:106-8.

Craib C.L. 1973. Breeding distribution of coastal cormorants. Witwatesrand Bird Club News Sheet, 82: 13.

Curry-Lindall K. 1970. Spread-wing postures in Pelecaniformes and Ciconiiformes. Auk 87:371-72.

Derenne P., G. Mary and Mougin J. L . 1976. Le cormoran à ventre blanc, Phalacrocorax albiventer melanogenis (Blyth) de l'archipel Crozet. Comité Natl. FranQaís Recherches Antarctiques 40:191-219.

Devillers P. & Terschuren J. A. 1978. Relationships between the blue-eyed shags of South America. Gerfaut, 68: 53-86.

Dorst J. & Mougin J. L . 1979. Order Pelecaniformes. Pp. 155-193, in: E. Mayr and G. W. Cottrell (eds.), Check List of Birds of the World, vol. L 2d ed. Museum of Comp. Zool., Cambridge, Mass.

Egremont P. & Rothschild M. 1979. The calculating cormorants. Biol. I. Linn. Soc., 12(2):181-86.

Elowson A. M. 1984. Spread-wing postures and the water repellency of feathers: a test of Rijke's hypothesis. Auk 101:371-83.

Fenwick C. D. & Browne W. M. M . 1975. Breeding of the spotted sbag at Whitewash Head, Banks Península. I. Roy. Soc. New Zealand 5:31-45.

Harris M. P. 1974. A complete census of the flightless cormorant (Nannopterum harrisi). Biol. Cons. 6:188-91.

Harris M. P. 1979. Population dynamics of the flightless cormorant Nannopterum harrisi. Ibis 121:135-46.

Hubbs C. L., A. L. Kelly and Limbaugh C . 1970. Diversity of feeding by Brandt's cormorant near San Diego. Calif. Fish & Game 56:156-65.

Huizinga H. W. 1971. Contracaeciasis in pelicaniform birds. J. Wildl. Dis., 7: 198-204.

Jones P. J. 1978. A possible function of the "wing-drying" posture in the Reed cormorant Phalacrocorax africanus. Ibis, 120: 540-542.

Kinsky F. C. 1970. "Spotted shags" in Westland. Notornis 17:102-4.

Lalas C. 1979. Double breeding season in pied shags on Stewart Island, from records by Roy Traill. Notornis, 26:94-95.

Miller B. 1979. Ecology of the little black cormorant Phalacrocorax sulcirostris and little pied cormorant P. melanoleucos in inland New South Wales. I. Food and feeding habits. Aust. Wildl. Res., 6:79~95.

Morrison M. L. & Slack R. D. 1977. The role of flock feeding in Olivaceus Cormorants. Bird Banding, 48: 277-279.

Morrison M. L. & Slack R. W. 1977. Population trends and status of the olivaceous cormorant. Amer. Birds 31:954-59.

Morrison, M. L., E. Shanley Jr. & Slack R. D. 1977. The food of nestling olivaceous cormorants. Southwest. Nat. 22:321-26.

Morrison M. L., R. D. Slack & Shanley E. 1978. Age and foraging ability relationships of Olivaceus Cormorants. Wilson Bull., 90 414-422.

Morrison, M. L., E. Shanley Jr. & Slack R. D. 1979. Breeding biology and age-specific mortality of olivaceous cormorants. Southwest. Nat., 24:259-66.

Nelson A. 1972. King shags in the MarIborough Sounds. Notornis, 18:30-37.

Norman F. 1974. Notes on the breeding of the pied cormorant near Werribee, Victoria, in 1971, 1972 and 1973. Emu, 74: 223-227.

Olrog C. C. 1975. Vagrancy of Neotropical cormorants, egrets and white ibis. Bird-Banding, 46: 207-12.

Olver M. D. & Kuyper M. A . 1978. Breeding biology of the Whitebreasted cormorant in Natal. Ostrich, 49: 25-30.

Potts K. J. 1977. Food of little shags and little black shags. Wildl. Rev., 8: 34-38.

Rijke A. M. 1970. The phylogenetic development of water repellency in water bird feathers. Ostrich, suppl., 8:67-76.

Robertson I. 1974. The food of nesting double-crested and pelagic cormorants at Mandarte Island, British Columbia, with notes on feeding ecology. Condor, 76: 346-348.

Saiff E. 1978. Anatomy of the middle ear of birds. The Pelecaniformes and Ciconiiformes. Zool. J. Linn. Soc. London, 63: 315-370.

SchIatter R. P. and Moreno C. A . 1976. Habitos alimentarios del cormorán Antartico, Phalacrocorax atriceps bransfieldensis (Murphy) en Isla Green, Antartico. Ser.Gent. Inst. Antarct. Chileno, 4:69-88.

Shaughnessy P.D. 1979. Further notes on Crowned cormorants on the coast of the South West Africa/Namibia. Cormorant, 6: 37-38.

Shaughnessy P.D. & Shaughnessy G.L. 1978. Crowned cormorants on the coast of the South West Africa/Namibia. Cormorant, 5: 21-25.

Siegfried W. R., A. J. Williams, P. C. H. Frost & Kinahan J. B. 1975. Plumage and ecology of cormorants. Zool. Afr., 10:183-192.

van Tets C. F. 1976. Further observations on the taking of pebbles by black-faced cormorants. Emu, 76:151-152.

van Tets C. F. 1976. Australasia and the origin of shags and cormorants, Phalacrocoracidae. Pp. 121-124, in: Proc. 16th Inter. Ornith. Congr., Canberra, Australia, 1974.

Voisin J. F. 1973. Notes on the Blue-eyed shags (genus Leucocarbo Bonaparte). Notornis, 20:262-271.

Watt J. P. C. 1975. Notes on Whero Island and other roosting and breeding stations of the Stewart Island shag (Leucocarbo carunculatus chalconotus). Notornis, 22:265-272.

Williams A. J. & Burger A. E. 1978. The ecology of the prey of Cape and bank cormorants. Cormorant, 4:28-29.

Williams A. J. & Burger A. E. 1979. Aspects of the breeding biology of the Imperial cormorant, Phalacrocorax atriceps, at Marion Island. Gerfaut, 69: 407-423.

1960 - 1969

Bartholomew G. A. Jr., R. C. Lasiewski & Crawford E. C. Jr. 1968. Patterns of panting and gular flutter in cormorants, pelicans, owls and doves. Condor, 70:31-34.

Bowen W., N. Gardiner, B. J. Harris & Thomas J. D. 1962. Communal nesting of Phalacrocorax africanus, Bubulcus ibis, and Anhinga rufa in southern Ghana. Ibis, 104:246-47.

Bowmaker A. P. 1963. Cormorant predation on two central African lakes. Ostrich, 34: 3-26.

Brodkorb P. 1963. Catalogue of fossil birds. Pt. I. Bull. Fla. State Mus. Biol. Sci., 7: 179-293.

Clark C. A. Jr. 1969. Spread-wing postures in Pelecaniformes, Ciconiiformes, and Falconiformes. Auk, 86:136-139.

Ford J. 1963. Dispersal and mortality in the pied cormorant in Western Australia. W. Aust. Nat., 8: 177-81.

Kury C. R. 1968. Difference in weight of male and female cormorants. Auk, 85: 513.

Lasiewski R. C. & Snyder C. K. 1969. Responses to high temperature in nestling double-crested and pelagic cormorants. Auk 86: 529-540.

Rand R. W. 1960. The biology of guano-producing sea-birds. 3. The distribution, abundance and feeding habits of the cormorants, Phalacrocoracidae, off the south west coast of the Cape Province. Invest. Rep. Div. Fish. Un. S. Afr., 42:5-32.

Rijke A. M. 1968. The water repellency and feather structure of cormorants, Phalacrocoracidae. Ostrich, 38:163-65.

Stonehouse B. 1967. Feeding behaviour and diving rhythms of some New Zeland Shags, Phalacrocoracidae. Ibis, 109: 600-605.

Snow B. 1966. Observations on the behaviour and ecology of the flightless cormorant Nannopterum harrisi. Ibis, 108: 265-280.

Thompson J. A. & Morley N. W. 1966. Physiological correlates of habitat selection in Australian cormorants. Emu, 66: 17-26.

van Tets G. F. 1965. A comparative study of some social communication patters in the Pelecaniformes. Am. Ornith. Union. Ornith. Monogr., No. 2.

van Tets G. F. 1968. White-breasted cormorant swallows pebbles on land. Emu, 67: 224.


Anonymous. 1949. Recovery round-up: little black cormorant. Aust. Bird Bander 14:55.

Behn F., J. D. Coodall, A. W. Johnson & Philippi R. A. 1955. The geographic distribution of the blue-eyed shags, Phalacrocorax albiventer and P. atriceps. Auk 72:6-13.

Bent A.C. 1922. Life Histories of North American Petrels and Pelicans and their allies. Dover Publications Inc. New York.

Du Plessis S.S. 1957. Growth and daily food intake of the White-breasted Cormorant in captivity. Ostrich, 28: 197-201.

Falla R. A. 1933. King shags of Queen Charlotte Sound. Emu 33:44-48.

Falla R. A. & Stokell G . 1945. Investigation of stomach contents of New Zealand fresh-water shags. Trans. Proc. Roy. Soc. N.Z. 74:320-31.

Harley K. L. S. 1946. Display and nesting of the littIe pied cormorant in the Brisbane Botanic Gardens. Emu 45:298-300.

Hoogerwerf A. 1953. [Phalacrocorax sulcirostris territori as a breeding-bird in West Java, 1952]. Limosa 26:71-79. (Dutch, English summary).

Godfrey R. 1932. The White-breasted cormorant. Blythswood Rev., 9: 82.

Kuroda N. 1925. On a new Japanese Cormorant, with some notes and suggestions. Tori, 4: 336­350. (Japanese with English summary).

Jordan R. 1959. El fenomeno de las regurgitaciones en el Guanay (Phalacrocorax bougainvillii L.) y un metodo para estimar la ingestion diaria. Bol. Cia. Adm. Guano, 35, 23-40.

Lanham U. N. 1947. Notes on the phylogeny of the Pelecaniformes. Auk, 64:65-70.

Laufer B. 1931. The domestication of the cormorant in China and Japan. Field Mus. Nat. Hist. (Anthropol. Ser.), 18:200-262.

Mack C. 1941. Cormorants and the Cippsland Lakes fishery. Mem. Nat. Mus. Vic., 12:95-117.

McKeown K. C. 1944. The food of cormorants and other fish-eating birds. Emu, 43:250-69.

McNally J. 1957. The feeding habits of cormorants in Victoria. Fauna Contr. Físh. & Game Dept. Vict., no. 6.

Meinertzhagen R. 1950. On the systematics of Phalacrocorax africanus Gmelin. Bull. Brit. Orn. Club, 70: 8.

Murphy R. C. 1916. Notes on American subantarctic cormorants. Bull. Am. Mus. Nat. Hist., 35:31-48.

PaIrngren P. 1935. Ein Exemplar von Phalacrocorax perspicillatus Pall. in den SammIungen des zoologischen Museums in Helsingfors. Ornis Fennica, 12:78-80.

Peters J. P. 1931. Check-list of the birds of the world. Vol. I. Harvard Univ. Press, Cambridge.

Rand R.W. 1960. The biology of guano-producing seabirds. 3. The distribution, abundance and feeding habitats of the cormorants Phalacrocoracidae off the southwest coast of the Cape Province. Investl. Rep. Div. Fish. Un. S. Afr., 42: 1-32.

Roberts W.W. 1934. Cape cormorant nesting in eastern cape Province. Blythswood Rev., 11: 34.

Serventy D. L. 1938. The feeding habits of cormorants in south-western Australia. Emu, 38:293-316.

Serventy D. L. 1939. Notes on cormorants. Emu, 38:357-71.

Shufeldt R. W. 1915. Comparative osteology of Harris' flight-less cormorant. Emu, 15:86-114.

Turbott E. G. 1956. Notes on the plumages and breeding cycle of the spotted shag Phalacrocorax (Stictocarbo) punctatus punctatus (Sparrman, 1786). Rec. Auckland (N. Z.) Inst., 4:343-363.

Williams L. 1942. Display & sexual behavior of the Brandt cormorant. Condor, 41:85-104.

Yamamoto H. 1967. [Phalacrocorax capillatus as a breeding bird on the Iwate Coast, Honshu.] Misc. Rept. Yamashina Inst. Ornith. Zool. 5:48-60. (Japanese, English summary).

Before 1900

Harting J. E. 1883. Fishing with Cormorants. Pp. 423-440 in Essays on sport and natural history. Horace Cox, London.

Ogilvie-Grant W.R. 1898. Steganopodes. pp. 329-484, in: Catalogue f the birds in the british Museum, vol. 26. British Museum (Nat. Hist.), London.

Stejneger L. 1889. Contribution to the history of the Pallas' cormorant. Proc. U. S. Natt. Mus., 12:83-88.

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