Editing Cormorant (section)

== Taxonomy ==
{{Update section|reason=several paragraphs ("The cormorant family are...", "Several evolutionary groups are...", and all but the last sentence of "In recent years, three...") appear to have been written in the mid-2000s and minimally updated since then, and as such are highly outdated, requiring extensive revision to reflect a modern state of knowledge of the relationships of cormorants and their relatives|date=June 2023}}
The genus ''Phalacrocorax'', from which the [[Family (taxonomy)|family]] name Phalacrocoracidae is derived, is [[Latin]]ised from [[Ancient Greek]] {{lang|grc|φαλακρός}} ''phalakros'' "bald" and {{lang|grc|κόραξ}} ''korax'' "raven".<ref name= job90>{{cite book | last= Jobling | first= James A. | year= 2010| title= The Helm Dictionary of Scientific Bird Names |url=https://archive.org/details/Helm_Dictionary_of_Scientific_Bird_Names_by_James_A._Jobling/page/n301 | publisher = Christopher Helm | location = London | isbn = 978-1-4081-2501-4 |page =301}}</ref> This is thought to refer to the ornamental white head plumes prominent in Mediterranean birds of this species, or the creamy white patch on the cheeks of adult [[great cormorant]]s, but is certainly not a unifying characteristic of cormorants.

The cormorant family was traditionally placed within the [[Pelecaniformes]] or, in the [[Sibley–Ahlquist taxonomy]] of the 1990s, the expanded [[Ciconiiformes]]. Pelecaniformes in the traditional sense—all waterbird groups with [[Bird feet and legs#Webbing and lobation|totipalmate]] foot webbing—are not a [[monophyletic]] group, even after the removal of the distantly-related [[tropicbird]]s. Their relationships and delimitation – apart from being part of a "higher waterfowl" [[clade]] which is similar but not identical to Sibley and Ahlquist's "pan-Ciconiiformes" – remain mostly unresolved. Notwithstanding, all evidence agrees that the cormorants and shags are closer to the [[darter]]s and [[Sulidae]] (gannets and boobies), and perhaps the pelicans or even [[penguin]]s, than to all other living birds.<ref>Kennedy ''et al.'' (2000), Mayr (2005)</ref>

In recent years, three preferred treatments of the cormorant family have emerged: either to leave all living cormorants in a single genus, ''Phalacrocorax'', or to split off a few species such as the [[imperial shag]] complex (in ''Leucocarbo'') and perhaps the [[flightless cormorant]]. Alternatively, the genus may be disassembled altogether and in the most extreme case be reduced to the [[Great cormorant|great]], [[White-breasted cormorant|white-breasted]] and [[Japanese cormorant]]s.<ref>See {{harvnb|Siegel-Causey|1988}}, Orta (1992) and Kennedy ''et al.'' (2000) for a review of classification schemes.</ref> In 2014, a landmark study proposed a 7 genera treatment, which was adopted by the [[IUCN Red List]] and [[BirdLife International]], and later by the [[International Ornithologists' Union|IOC]] in 2021, standardizing it.<ref name=IOU/><ref name=kennedy2014/>

[[File:Xiphoid_phalacrocorax.jpg|thumb|Occipital crest or ''os nuchale'' in ''Phalacrocorax carbo'']]
The cormorants and the darters have a unique bone on the back of the top of the skull known as the ''os nuchale'' or occipital style which was called a xiphoid process in early literature. This bony projection provides anchorage for the muscles that increase the force with which the lower mandible is closed.<ref name=Yarrell>{{cite journal|title=On the xiphoid bone and its muscles in the Corvorant (Pelecanus carbo) |author=Yarrell, William |pages= 234–237|url=https://www.biodiversitylibrary.org/page/2310645| journal=The Zoological Journal| volume=4 |year= 1828}}</ref><ref>{{cite journal|doi=10.1111/j.1096-3642.1876.tb02572.x|title=1. Notes on the Anatomy of Plotus anhinga|journal=Proceedings of the Zoological Society of London|volume=44|pages=335–345|year=2009|last1=Garrod|first1=A. H.|url=https://www.biodiversitylibrary.org/part/73591}}</ref> This bone and the highly developed muscles over it, the M. adductor mandibulae caput nuchale, are unique to the families Phalacrocoracidae and Anhingidae.<ref>{{cite journal|doi=10.1080/00445096.1978.11447608|title=Functional Anatomy of the Feeding Apparatus of Four South African Cormorants|journal=Zoologica Africana|volume=13|pages=81–102|year=2015|last1=Burger|first1=A E|doi-access=free}}</ref><ref>{{cite journal|doi=10.1071/MU915086|title=Comparative osteology of Harris's Flightless Cormorant (Nannopterum harrisi)|journal=Emu|volume=15|issue=2|pages=86–114|year=1915|last1=Shufeldt|first1=R.W.|bibcode=1915EmuAO..15...86S |url=https://zenodo.org/record/1944350}}</ref>

Several [[evolution]]ary groups are still recognizable. However, combining the available evidence suggests that there has also been a great deal of [[convergent evolution]]; for example, the cliff shags are a convergent [[paraphyletic]] group. The proposed division into ''Phalacrocorax sensu stricto'' (or [[subfamily]] "Phalacrocoracinae") cormorants and ''Leucocarbo sensu lato'' (or "Leucocarboninae") shags{{sfn|van Tets|1976}}{{sfn|Siegel-Causey|1988}} does have some degree of merit.<ref name = kennedyetal2000>Kennedy ''et al.'' (2000)</ref> The resolution provided by the [[mtDNA]] [[12S rRNA]] and [[ATPase]] [[Protein subunit|subunits]] six and eight [[DNA sequence|sequence]] data<ref name = kennedyetal2000 /> is not sufficient to resolve several groups to satisfaction properly; in addition, many species remain unsampled, the fossil record has not been integrated in the data, and the effects of hybridisation – known in some Pacific species especially – on the DNA sequence data are unstudied.

A multigene [[molecular phylogenetic]] study published in 2014 provided a genus-level phylogeny of the family;<ref name=kennedy2014>{{Cite journal | last1=Kennedy | first1=M. | last2=Spencer | first2=H.G. | date=2014 | title=Classification of the cormorants of the world | journal=Molecular Phylogenetics and Evolution | volume=79 | pages=249–257 | doi=10.1016/j.ympev.2014.06.020| pmid=24994028 | bibcode=2014MolPE..79..249K }}</ref> this is now followed by most authorities, including the [[IOC World Bird List]].<ref name="IOU"/>

{{Clade | style=font-size:100%;line-height:90%
|label1=Phalacrocoracidae
|1={{Clade
   |1=''[[Microcarbo]]'' – Five species
   |2={{clade
      |1=''[[Poikilocarbo]]'' – Red-legged cormorant
      |2={{clade
         |1={{clade
            |1=''[[Urile]]'' – Four species (one extinct)
            |2=''[[Phalacrocorax]]'' – Eleven species
            }}
         |2={{clade
            |1=''[[Gulosus]]'' – European shag
            |2={{clade
               |1=''[[Nannopterum]]'' – Three species
               |2=''[[Leucocarbo]]'' – 15 species
               }}
            }}
         }}
      }}
   }}
}}

===List of genera===
{{Main|List of cormorants}}
[[File:Cormorant diving for food in Morro Bay.jpg|thumb|Cormorant (species unknown) begins its dive]]
[[File:Cormoran Shag.jpg|thumb|Immature [[imperial shag]] (''Leucocarbo atriceps'')]]
[[File:Little Cormorant (Phalacrocorax niger) in Hyderabad W IMG 8389.jpg|thumb|Little cormorant ''(Microcarbo niger)'' in [[Hyderabad, India]]]]
[[File:Guanokormoran (Phalacrocorax bougainvillii) - Weltvogelpark Walsrode 2012-01.jpg|thumb|[[Guanay cormorant]] (''Leucocarbo bougainvillii'') at [[Weltvogelpark Walsrode]]]]
[[File:Japanese cormorant - kanagawa japan - 2024 feb 25.webm|thumb|[[Japanese cormorant]] in [[Kanagawa Prefecture|Kanagawa]], [[Japan]]]]
As per the [[International Ornithologists' Union|IOU]], the [[IUCN Red List]] and [[BirdLife International]], the family contains 7 genera:<ref name="IOU">{{cite web | title=Storks, frigatebirds, boobies, darters, cormorants – IOC World Bird List | website=IOC World Bird List – Version 15.1 | date=2025-02-20 | url=https://www.worldbirdnames.org/new/bow/storks/ | access-date=2025-04-14}}</ref>

{| class="wikitable"
|-
! Image !! Genus  !! Species 
|-
|[[File:Little cormorant (Microcarbo niger) - 20070322.jpg|175px]] || ''[[Microcarbo]]'' {{small|Bonaparte, 1856}} || 
*[[Crowned cormorant]] ''Microcarbo coronatus''
*[[Little cormorant]] ''Microcarbo niger''
*[[Little pied cormorant]] ''Microcarbo melanoleucos''
*[[Pygmy cormorant]] ''Microcarbo pygmeus''
*[[Reed cormorant]] or long-tailed cormorant ''Microcarbo africanus''
*†Serventys' cormorant ''[[Microcarbo serventyorum]]''
Around Indian Ocean, one species extending from Central Asia into Europe. Mostly in freshwater habitat. Small (about 50–60&nbsp;cm long), nondescript black to dark brown (except for one species with white underparts).
|-
|[[File:Phalacrocorax gaimardi 00.jpg|175px]] || ''[[Poikilocarbo]]'' {{small|Boetticher, 1935}} || 
*[[Red-legged cormorant]] ''Poikilocarbo gaimardi''
Subtropical to subantarctic Pacific South America, ranging a bit into the southwestern Atlantic. Maritime. Mid-sized (around 75&nbsp;cm), grey with scalloped wings and contrasting white/yellow/red neck mark and bare parts. Its high-pitched chirping calls are quite unlike those of other cormorants.
|-
|[[File:Red-faced Cormorant on Pribilof Islands, 5-1979 2.jpg|175px]] || ''[[Urile]]'' {{small|Bonaparte, 1855}} || 
*[[Brandt's cormorant]] ''Urile penicillatus''
*[[Red-faced cormorant]] ''Urile urile''
*[[Pelagic cormorant]] ''Urile pelagicus''
*[[Spectacled cormorant]] †''Urile perspicillatus''
Northern Pacific, one species extending into subtropical waters on the American West Coast. Maritime. Smallish to large (65–100&nbsp;cm), generally black with metallic sheen (usually blue/green), in breeding plumage with bright bare facial skin in the eye region and two crests (crown and nape).
|-
|[[File:Great cormorants at the Tama river.JPG|175px]] || ''[[Phalacrocorax]]'' {{small|Brisson, 1760}} ||
*[[Bank cormorant]]'' Phalacrocorax neglectus''
*[[Socotra cormorant]] ''Phalacrocorax nigrogularis''
*[[Pitt shag]]'' Phalacrocorax featherstoni''
*[[Spotted shag]] ''Phalacrocorax punctatus''
*[[Black-faced cormorant]] ''Phalacrocorax fuscescens''
*[[Australian pied cormorant]] ''Phalacrocorax varius''
*[[Little black cormorant]] ''Phalacrocorax sulcirostris''
*[[Indian cormorant]] ''Phalacrocorax fuscicollis''
*[[Cape cormorant]] ''Phalacrocorax capensis''
*[[Japanese cormorant]] or Temminck's cormorant ''Phalacrocorax capillatus''
*[[Great cormorant]] (including [[white-breasted cormorant]] and black shag) ''Phalacrocorax carbo''
Mostly around the Indian Ocean, one species group extending throughout Eurasia and to Atlantic North America. Maritime to freshwater. Size very variable (60–100&nbsp;cm), blackish with metallic sheen (usually bronze to purple) and/or white cheek and thigh patches or underside at least in breeding plumage; usually a patch of bare yellow skin at the base of the bill.
|-
|[[File:Shag, Bangor - geograph.org.uk - 1243477.jpg|175px]] || ''[[European shag|Gulosus]]'' {{small|Montagu, 1813}} || 
*[[European shag]] ''Gulosus aristotelis''
Breeds in the European Arctic, winters in Europe and North Africa. Maritime. Mid-sized (70–80&nbsp;cm), glossy black, in breeding plumage with a forehead crest curled to the front.
|-
|[[File:Double-crested Cormorant (49605889903).jpg|175px]] || ''[[Nannopterum]]'' {{small|Sharpe, 1899}} || 
*[[Flightless cormorant]] ''Nannopterum harrisi''
*[[Neotropic cormorant]] ''Nannopterum brasilianum''
*[[Double-crested cormorant]] ''Nannopterum auritum''
Throughout the Americas. Mostly freshwater. Smallish to large (65–100&nbsp;cm), nondescript brownish-black. One species with white tufts on sides of head in breeding plumage.
|-
|[[File:Phalacrocorax atriceps, Beagle Channel.jpg|175px]] || ''[[Blue-eyed shag|Leucocarbo]]'' {{small|Bonaparte, 1856}} || 
*[[Rock shag]] or Magellanic cormorant ''Leucocarbo magellanicus''
*[[Guanay cormorant]] ''Leucocarbo bougainvillii''
*[[Bounty shag]] ''Leucocarbo ranfurlyi''
*[[New Zealand king shag]] or rough-faced shag ''Leucocarbo carunculatus ''
*[[Chatham shag]] ''Leucocarbo onslowi''
*[[Stewart Island shag]] (including [[Foveaux shag]] ''L. stewarti'') ''Leucocarbo chalconotus''
*[[Auckland shag]] ''Leucocarbo colensoi''
*[[Campbell shag]] ''Leucocarbo campbelli''
*[[Imperial shag]] or blue-eyed shag ''Leucocarbo atriceps''
*[[South Georgia shag]] ''Leucocarbo georgianus''
*[[Crozet shag]] ''Leucocarbo melanogenis''
*[[Antarctic shag]] ''Leucocarbo bransfieldensis''
*[[Kerguelen shag]] ''Leucocarbo verrucosus''
*[[Heard Island shag]] ''Leucocarbo nivalis''
*[[Macquarie shag]] ''Leucocarbo purpurascens''
Generally Subantarctic, but extending farther north in South America; many oceanic-island endemics. Maritime. Smallish to largish (65–80&nbsp;cm), typically black above and white below, and with bare yellow or red skin in the facial region. A southern circumpolar group of several species (the blue-eyed shag complex) is characterised by bright blue orbital skin.
|-
|}

Prior to 2021, the IOU (or formerly the IOC) classified all these species in just three genera: ''Microcarbo'', ''Leucocarbo'', and a broad ''Phalacrocorax'' containing all remaining species; however, this treatment rendered ''Phalacrocorax'' deeply paraphyletic with respect to ''Leucocarbo''. Other authorities, such as the [[The Clements Checklist of Birds of the World|Clements Checklist]], formerly recognised only ''Microcarbo'' as a separate genus from ''Phalacrocorax''.

===Evolution and fossil record===
{{More citations needed section|date=November 2024}}
The details of the evolution of the cormorants are mostly unknown. Even the technique of using the distribution and relationships of a species to figure out where it came from, biogeography, usually very informative, does not give very specific data for this probably rather ancient and widespread group. However, the closest living relatives of the cormorants and shags are the other families of the [[suborder]] [[Sulae]]—[[darter]]s and [[Sulidae|gannets and boobies]]—which have a primarily [[Gondwana]]n distribution. Hence, at least the modern diversity of Sulae probably originated in the southern hemisphere.

While the Leucocarbonines are almost certainly of southern Pacific origin—possibly even the Antarctic which, at the time when cormorants evolved, was not yet ice-covered—all that can be said about the Phalacrocoracines is that they are most diverse in the regions bordering the Indian Ocean, but generally occur over a large area.

Similarly, the origin of the family is shrouded in uncertainties. Some [[Late Cretaceous]] fossils have been proposed to belong with the Phalacrocoracidae:<br />
A [[scapula]] from the [[Campanian]]-[[Maastrichtian]] boundary, about 70 mya (million years ago), was found in the [[Nemegt Formation]] in Mongolia; it is now in the [[Paleontological Institute of Russian Academy of Sciences|PIN]] collection.<ref>Kurochkin (1995)</ref> It is from a bird roughly the size of a spectacled cormorant, and quite similar to the corresponding bone in ''Phalacrocorax''. A [[Maastrichtian]] (Late Cretaceous, c. 66 mya) right [[femur]], [[AMNH]] FR 25272 from the [[Lance Formation]] near [[Lance Creek, Wyoming]], is sometimes suggested to be the second-oldest record of the Phalacrocoracidae; this was from a rather smaller bird, about the size of a [[long-tailed cormorant]].<ref>Hope (2002)</ref> However, cormorants likely originated much later, and these are likely misidentifications.<ref name="Kuhl Frankl-Vilches Bakker et al Unbiased Molecular Approach">{{cite journal |last1=Kuhl |first1=Heiner |last2=Frankl-Vilches |first2=Carolina |last3=Bakker |first3=Antje |last4=Mayr |first4=Gerald |last5=Nikolaus |first5=Gerhard |last6=Boerno |first6=Stefan T |last7=Klages |first7=Sven |last8=Timmermann |first8=Bernd |last9=Gahr |first9=Manfred |title=An Unbiased Molecular Approach Using 3′-UTRs Resolves the Avian Family-Level Tree of Life |journal=Molecular Biology and Evolution |date=4 January 2021 |volume=38 |issue=1 |pages=108–127 |doi=10.1093/molbev/msaa191 |pmid=32781465 |pmc=7783168 |doi-access=free}}</ref>

As the [[Early Oligocene]] ''"Sula" ronzoni'' cannot be assigned to any of the sulid families—cormorants and shags, darters, and gannets and boobies—with certainty, the best interpretation is that the Phalacrocoracidae diverged from their closest ancestors in the Early Oligocene, perhaps some 30 million years ago, and that the Cretaceous fossils represent ancestral sulids, "pelecaniforms" or "higher waterbirds"; at least the last lineage is generally believed to have been already distinct and undergoing [[evolutionary radiation]] at the [[Cretaceous–Paleogene extinction event|end of the Cretaceous]]. What can be said with near certainty is that AMNH FR 25272 is from a diving bird that used its feet for underwater locomotion; as this is liable to result in some degree of convergent evolution and the bone is missing indisputable neornithine features, it is not entirely certain that the bone is correctly referred to this group.<ref>Hope (2002) and see [[Hesperornithes]]</ref>

Phylogenetic evidence indicates that the cormorants diverged from their closest relatives, the darters, during the Late Oligocene, indicating that most of the claims of Cretaceous or early Paleogene cormorant occurrences are likely misidentifications.<ref name="Kuhl Frankl-Vilches Bakker et al Unbiased Molecular Approach"/>

During the late Paleogene, when the family presumably originated, much of Eurasia was covered by shallow seas, as the [[Indian Plate]] finally attached to the mainland. Lacking a detailed study, it may well be that the first "modern" cormorants were small species from eastern, south-eastern or southern Asia, possibly living in freshwater habitat, that dispersed due to [[tectonics|tectonic]] events. Such a scenario would account for the present-day distribution of cormorants and shags and is not contradicted by the fossil record; as remarked above, a thorough review of the problem is not yet available.

[[File:Doublecrestcorm14.jpg|thumb|upright|Double-crested cormorant]]

Even when ''Phalacrocorax'' was used to unite all living species, two distinct genera of prehistoric cormorants became widely accepted today:
* ''[[Limicorallus]]'' (Indricotherium middle Oligocene of Chelkar-Teniz, Kazakhstan)<!-- BullAMNH151:1. -->
* ''[[Nectornis]]'' (Late Oligocene/Early Miocene of Central Europe – Middle Miocene of Bes-Konak, Turkey) – includes ''Oligocorax miocaenus''<!-- AnnNHMusWien104A:237; Auk52:75; Condor58:367; Geobios36:719 -->

The proposed genus ''Oligocorax'' appears to be [[paraphyletic]]; the European species have been separated in ''Nectornis'', and the North American ones placed in the expanded ''Phalacrocorax''; the latter might just as well be included in ''Nannopterum''. A [[Late Oligocene]] fossil cormorant foot from [[Enspel]], Germany, sometimes placed in ''Oligocorax''<!-- AnnNHMusWien104A:237 -->, would then be referable to ''Nectornis'' if it proves not to be too distinct. ''Limicorallus'', meanwhile, was initially believed to be a [[rail (bird)|rail]] or a [[dabbling duck]] by some. There are also undescribed remains of apparent cormorants from the [[Quercy Phosphorites]] of [[Quercy]] (France), dating to some time between the [[Late Eocene]] and the mid-[[Oligocene]]. All these early European species might belong to the basal group of "microcormorants", as they conform with them in size and seem to have inhabited the same habitat: subtropical coastal or inland waters. While this need not be more than [[convergent evolution|convergence]], the phylogeny of the modern (sub)genus ''Microcarbo'' &ndash; namely, whether the Western Eurasian ''M. pygmaeus'' is a [[basal (evolution)|basal]] or highly derived member of its clade &ndash; is still not well understood at all as of 2022.

Some other Paleogene remains are sometimes assigned to the Phalacrocoracidae, but these birds seem rather intermediate between cormorants and darters (and lack clear [[autapomorph]]ies of either). Thus, they may be quite basal members of the [[Palacrocoracoidea]]. The [[taxa]] in question are:
* ''[[Piscator (bird)|Piscator]]'' (Late Eocene of England)
* "Pelecaniformes" gen. et sp. indet. (Jebel Qatrani Early Oligocene of Fayum, Egypt) – similar to ''Piscator''?
* ''[[Borvocarbo]]'' (Late Oligocene of C Europe)

The supposed Late Pliocene/Early Pleistocene ''"[[Valenticarbo]]"'' is a ''[[nomen dubium]]'' and given its recent age probably not a separate genus.
[[File:Phalacrocorax filyawi FLMNH.jpg|thumb|Reconstructed fossil skeleton of the extinct [[Miocene]]-aged ''[[Phalacrocorax filyawi]]'', [[Florida Museum of Natural History]]]]
The remaining fossil species are not usually placed in a modern phylogenetic framework. While the numerous western US species are most likely prehistoric representatives of the coastal ''Urile'' or inland ''Nannopterum'', the European fossils pose much more of a problem due to the singular [[common shag]] being intermediate in size between the other two European cormorant lineages, and as of 2022 still of [[ghost lineage|mysterious ancestry]]; notably, a presumably lost collection of Late Miocene fossils from the [[Odesa]] region may have contained remains of all three (sub)genera inhabiting Europe today. Similarly, the Plio-Pleistocene fossils from Florida have been allied with ''Nannopterum'' and even ''Urile'', but may conceivably be ''Phalacrocorax''; they are in serious need of revision since it is not even clear how many species are involved. Provisionally, the fossil species are thus all placed in ''Phalacrocorax'' here:
* ''Phalacrocorax marinavis'' (Oligocene – Early Miocene of Oregon, US) &ndash; formerly ''Oligocorax''; ''Urile'' or ''Nannopterum''?<!-- Condor58:367; ContribMusPaleontolUnivMich24:205 -->
* ''Phalacrocorax littoralis'' (Late Oligocene/Early Miocene of St-Gérand-le-Puy, France) &ndash; formerly ''Oligocorax''; ''Nectornis''?<!-- AnnNHMusWien104A:237; Auk52:75; Condor58:367; Geobios36:719 -->
* ''Phalacrocorax intermedius'' (Early – Middle Miocene of C Europe) &ndash; includes ''P. praecarbo, Ardea/P. brunhuberi'' and ''Botaurites avitus''; ''Microcarbo'', ''Phalacrocorax'' or ''Gulosus''?<!-- AnnNHMusWien104A:237;Condor58:367 -->
* ''Phalacrocorax macropus'' (Early Miocene – Pliocene of north-west US) &ndash; ''Urile'' or ''Nannopterum''?<!-- (Middle) Pleistocene?! Auk30:29; Condor58:367; Condor69:24; ContribMusPaleontolUnivMich24:205 -->
* ''Phalacrocorax ibericus'' (Late Miocene of Valles de Fuentiduena, Spain) &ndash; ''Microcarbo'', ''Phalacrocorax'' or ''Gulosus''?<!-- AnnNHMusWien104A:237 -->
* ''Phalacrocorax lautus'' (Late Miocene of Golboçica, Moldavia) &ndash; ''Microcarbo'', ''Phalacrocorax'' or ''Gulosus''?<!-- AnnNHMusWien104A:237 -->
* ''Phalacrocorax serdicensis'' (Late Miocene of Hrabarsko, Bulgaria); ''Microcarbo'', ''Phalacrocorax'' or ''Gulosus''?<!-- AnnNHMusWien104A:237 -->
* ''Phalacrocorax'' sp(p). (Late Miocene of Odesa region, Ukraine) &ndash; up to 4 species, one of which is probably ''P. longipes''; ''Microcarbo'', ''Phalacrocorax'' and/or ''Gulosus''?<ref>These are the fossils described in the same paper as ''[[Pelecanus odessanus]]'' and sometimes cited as "''Phalacrocorax'' sp. Wildhalm", which is a widespread ''[[lapsus calami]]'' or printing error for the initial describer, Ignatiy Vidgal'm (Игнатий Видгальм in Russian, a German emigrant originally named Ignaz Wi(e)dhalm and often transcribed as "J. Widhalm").<!-- Born in Regensburg, emigrated to Odessa. Second surname is usually given as "D.", but also "Martynovich"; the latter is probably a Russified patronymic, the former his original German second surname. See http://odessa-memory.info/index.php?id=507. Transcription in Olson 1999 is somewhat erroneous. "Ignaz/Ignatiy" was probably abbreviated as "J." by Wi(e)dhalm himself because this letter was often used interchangeably with capital "I" in contemporary German. The German family name was properly "Wiedhalm", but the "e" is often dropped even in the German version (see e.g. "Leopold Widhalm" article), and in any case does not carry over to the Russian version. The error "Wildhalm" is attested as early as 1916 (contra Olson 1999), in Lambrecht's  papers "Die Gattung Plotus im ungarischen Neogen" and "A madarak palaeontologiájának története és irodalma"; it may have been due to a damaged letter "E", as author names were usually typeset in small caps at that time. --> He discussed three presumed species of cormorant under the provisional non-[[Linnean nomenclature|Linnean]] names "Haliaeus fossilis, var. Odessana major, medius, [a]nd minor" ("fossil cormorant, large/mid-sized/small Odessan variety").<!-- See Olson 1999: A new species of pelican (Aves: Pelecanidae) from the Lower Pliocene of North Carolina and Florida https://repository.si.edu/bitstream/handle/10088/6492/VZ_293_Pelecanus_schreiberi.pdf?sequence=1&isAllowed=y --> While various different bones were assigned to the large species (and eventually referred to ''P. longipes''), one small and one mid-sized tarsometatarsus fragment remained unassigned; a few proximal ends of the same bone were provisionally included in the large species, but exceed its more complete tarsometatarsi in size and may represent a distinct and even larger fourth species. The fossils are probably lost nowadays and even the original publication is held by very few libraries; this has so far prevented a thorough review of the remains, but one partial [[coracoid]] does not appear to belong to ''Phalacrocorax [[sensu stricto]]'' and may have been closer in [[Morphology (biology)|habitus]] to North Pacific shags (''Urile''), but is unlikely to have been closely related{{Verify source|date=November 2007}} to these: Howard (1932).</ref>
* ''Phalacrocorax femoralis'' (Modelo Late Miocene/Early Pliocene of WC North America) &ndash; formerly ''Miocorax''; ''Nannopterum''?<!-- Auk52:75; Condor31:167; Condor58:367 -->
* ''Phalacrocorax'' sp. (Late Miocene/Early Pliocene of Lee Creek Mine, US) &ndash; ''Nannopterum'' or ''Phalacrocorax''?
* ''Phalacrocorax'' sp. 1 (Late Miocene/Early Pliocene of WC South America) &ndash; probably ''Leucocarbo''<!-- "Phalacrocorax aff.bougainvillii" -->
* ''Phalacrocorax'' sp. 2 (Pisco Late Miocene/Early Pliocene of SW Peru) &ndash; ''Poikilocarbo'' or ''Leucocarbo''?
* ''Phalacrocorax longipes'' (Late Miocene – Early Pliocene of Ukraine) &ndash; formerly ''Pliocarbo''; ''Microcarbo'', ''Phalacrocorax'' or ''Gulosus''?<!-- AnnNHMusWien104A:237 -->
* ''Phalacrocorax goletensis'' (Early Pliocene – Early Pleistocene of Mexico) &ndash; ''Urile'' or ''Nannopterum'', perhaps ''Poikilocarbo'' or ''Leuocarbo''<!-- ContribMusPaleontolUnivMich24:205 -->
* ''Phalacrocorax wetmorei'' (Bone Valley Early Pliocene of Florida) &ndash; ''Nannopterum'' or ''Phalacrocorax''?<!-- Condor58:367; Condor72:293 -->
* ''Phalacrocorax'' sp. (Bone Valley Early Pliocene of Polk County, Florida, US) &ndash; ''Nannopterum'' or ''Phalacrocorax''?<ref>A [[Anatomical terms of location#Proximal and distal|proximal]] [[ulna]], Specimen PB 311, [[Pierce Brodkorb]] collection. Initially assigned to ''P. idahensis''. However, it is far too large, being from a very big species possibly larger than a great cormorant: Murray (1970).</ref>
* ''Phalacrocorax leptopus'' (Juntura Early/Middle Pliocene of [[Juntura, Oregon|Juntura]], [[Malheur County, Oregon|Malheur County]], Oregon, US) &ndash; ''Nannopterum''?
* ''Phalacrocorax reliquus'' (Middle Pliocene of Mongolia) &ndash; ''Microcarbo'', ''Phalacrocorax'' or ''Gulosus''?
* ''Phalacrocorax idahensis'' (Middle Pliocene – Pleistocene of Idaho, US, and possibly Florida) &ndash; ''Nannopterum''?<!-- Condor58:367; Condor72:293; ContribMusPaleontolUnivMich24:205 -->
* ''Phalacrocorax destefanii''{{Verify source|date=November 2007}}<!-- destefani? --> (Late Pliocene of Italy) &ndash; formerly ''Paracorax''; ''Microcarbo'', ''Phalacrocorax'' or ''Gulosus''?<!-- Condor58:367 -->
* ''Phalacrocorax filyawi'' (Pinecrest Late Pliocene of Florida, US) &ndash; may be ''P. idahensis''; ''Nannopterum'' or ''Phalacrocorax'', perhaps ''Urile''?
* ''Phalacrocorax kennelli'' (San Diego Late Pliocene of California, US) &ndash; ''Urile'' or ''Nannopterum''?<!-- Condor72:293 -->
* ''Phalacrocorax kumeyaay'' (San Diego Late Pliocene of California, US) &ndash; ''Urile'' or ''Nannopterum''?<!-- OccasionalPapersOfTheUniversityOfKansasMuseumOfNaturalHistory140:1. -->
* ''Phalacrocorax macer'' (Late Pliocene of Idaho, US) &ndash; ''Nannopterum''?<!-- Condor72:293 -->
* ''Phalacrocorax mongoliensis'' (Late Pliocene of W Mongolia) &ndash; ''Microcarbo'', ''Phalacrocorax'' or ''Gulosus''?
* ''Phalacrocorax'' sp. (La Portada Late Pliocene of N Chile) &ndash; may be same as Late Miocene/Early Pliocene "''Phalacrocorax'' sp. 2"; ''Poikilocarbo'' or ''Leucocarbo''?
* ''Phalacrocorax rogersi'' (Late Pliocene – Early Pleistocene of California, US) &ndash; ''Urile'' or ''Nannopterum''?<!-- Condor33:31; Condor34:118; Condor58:367; ContribMusPaleontolUnivMich24:205 -->
* ''Phalacrocorax chapalensis'' (Late Pliocene/Early Pleistocene of Jalisco, Mexico) &ndash; ''Urile'' or ''Nannopterum'', perhaps ''Poikilocarbo'' or ''Leucocarbo''?<!-- *ContribMusPaleontolUnivMich24:205 -->
* ''Phalacrocorax gregorii'' (Late Pleistocene of Australia) &ndash; possibly not a valid species; ''Microcarbo'', ''Phalacrocorax'' or ''Leucocarbo''?<!-- Condor58:367 -->
* ''Phalacrocorax vetustus'' (Late Pleistocene of Australia) &ndash; formerly ''Australocorax'', possibly not a valid species; ''Microcarbo'', ''Phalacrocorax'' or ''Leucocarbo''?<!-- Condor58:367 -->
* ''Phalacrocorax'' sp. (Sarasota County, Florida, US) &ndash; may be ''P. filawyi/idahensis''; ''Nannopterum'' or ''Phalacrocorax''?<!-- Plio- or Plestocene: Forsberg  Shell  Pit, Leisey Shell Pit, Macasphalt Shell Pit or Richardson Road Shell Pit? -->

The former ''"Phalacrocorax"'' (or ''"Oligocorax"'') ''mediterraneus'' is now considered to belong to the [[Bathornithidae|bathornithid]] ''[[Paracrax antiqua]]''.<ref>Cracraft (1971)</ref> ''"P." subvolans'' was actually a darter (''Anhinga'').