Belongs within: Aequornithia.
Contains: Phalacrocoracidae.
The Sulida or Suloidea are the clade including Sulidae (gannets and boobies), Anhingidae (darters) and Phalacrocoracidae (shags or cormorants). All members of this clade are diving fish-eating birds.
The darters or anhingas of the Anhingidae, all living species of which are placed in the single genus Anhinga, are a pantropical group of birds with particularly long and flexible necks; the neck is held back against the body when approaching prey, then darted forward to impale the prey on their long straight beak. The late Oligocene Oligocrax stoeffelensis resembled a small cormorant in overall appearance and may have represented either a stem-group phalacrocoracid or a stem member of the Phalacrocoracidae + Anhingidae clade (Mayr 2009). Borvocarbo guilloti is another late Oligocene species based on a coracoid with a plesiomorphic scapular articulation relative to Phalacrocoracidae and Anhingidae (Mayr 2009).
The Sulidae contain the gannets and boobies, marine birds that capture fish by plunge diving. Sulids nest in colonies, sometimes very densely. The gannets of the genus Morus are found in temperate waters of the Northern and Southern Hemispheres whereas the boobies of the genus Sula are tropical in distribution. The Morus species are all mostly white with black-tipped wings and buff skullcaps. Abbott’s booby Papasula abbotti is an endangered species endemic to Christmas Island in the eastern Indian Ocean.
Boobies
Published 27 January 2009
The Sulidae are the boobies and gannets, a small but fairly cosmopolitan assortment of seabirds. Among the living sulids, the gannets form the genus Morus, and most boobies belong to the genus Sula. The exception is Abbott’s booby Papasula abbotti which was originally included in Sula, and is still commonly referred to as such (at least in popular sources), but sits on the gannet rather than the booby side of the divergence between the two main genera (Friesen & Anderson 1997). The distinctions between the three genera are not huge, and some authors in the past have referred to all living sulids as Sula. In the recent fauna there is a clear geographical division between gannets and boobies—Morus is found in the northern and southern temperate zones while Sula and Papasula are tropical or subtropical—but this does not appear to have always been so in the past. The Pliocene Pisco Formation of Peru has provided species assigned to both Sula and Morus (Stucchi & Urbina 2004).
Eight fossil genera have been assigned to Sulidae (as well as fossil species of Sula and Morus*)—the Eocene Masillastega and Eostega, the Oligocene Empheresula, the Miocene Microsula (now a synonym of Morus), Miosula, Enkurosula and Sarmatosula, and the Pliocene Palaeosula and Ramphastosula. The Cretaceous Elopteryx did a stint as a close relative of the sulids, but is now regarded as a dinosaur of the Troodontidae and not even a bird (for most definitions of the word “bird”). Eostega lebedinskyi is known from a single mandible that was recently redescribed by Mlíkovský (2007), who reasserted its sulid nature (past authors have disagreed). Masillastega rectirostris is known from a skull from the famed Messel formation, and differs from living sulids in having a comparatively long beak (Mayr 2002). It is also distinct in having seemingly inhabited a freshwater environment, while all modern species are exclusively marine. Mayr (2002) only tentatively regarded Masillastega as a sulid, and it may be a stem-member of the family (Mayr 2005). Mlíkovský (2007) synonymised the two Eocene genera on the basis of a lack of significant differences between them.
*Apart from a single taxon described as a subspecies of the modern species (Papasula abbotti costelloi [ha ha]), there don’t appear to be any fossil species assigned to Papasula. Considering the only recent distinction of Papasula from Sula, one wonders whether any fossils of the former are masquerading as the latter.
Empheresula arvernensis is represented by a pelvis from France (the original material also included a sternum, but Lambrecht later indicated that the sternum was not even sulid—Mlíkovský 2002). The French Oligocene also provided Sula ronzoni, which, if correctly assigned (which seems to be debatable*), would suggest that the living sulid lineages had diverged by that point (Friesen & Anderson 1997, used a molecular clock to estimate a divergence time for Sula vs. Morus/Papasula of 23 million years ago, which is also consistent with this, but the age calculation methods used by Friesen & Anderson can only be described as [ahem] dated). The European Miocene genera Enkurosula and Sarmatosula are both known from isolated humeri, and are both doubtfully distinct from Morus (Nelson 2006; Olson, 1984, suggests that Microsula (=Enkurosula) pygmaea may be conspecific with the contemporaneous Microsula avita of Maryland in the United States, which has itself been since reassigned to Morus), as are the Californian genera Miosula and Palaeosula.
*I’m rather confused here. Nelson (2006) notes that Sula ronzoni has four notches on the sternum, and indicates that this would place it on the sulid stem. However, Mlíkovský (2002, 2007) states that the type material of S. ronzoni is an incomplete pelvis, so what is Nelson talking about?
The wierdest of all sulids, though, is the Pliocene Peruvian Pisco Formation’s Ramphastosula ramirezi. The genus name means “toucan-booby” and is undeniably appropriate as Ramphastosula, instead of having a dagger-like straight beak like all other sulids, had a deep beak with a distinct arch as shown in the reconstruction above from Stucchi & Urbina (2004). The skull of Ramphastosula is also more robust than in other sulids, seemingly to support the enlarged beak. Ramphastosula was obviously pursuing a different lifestyle to other sulids, as it looks as if it would be ill-suited to catching fish by plunge-diving. Stucchi & Urbina (2004) suggest that its robust skull indicates greater diving ability than other sulids, so perhaps Ramphastosula was more inclined to pursue its prey underwater than its modern relatives. Unfortunately, no post-cranial material is known as yet for this species.
Systematics of Sulida
Synapomorphies (Mayr 2004): Os palatinum a flat horizontal plate; vomer absent; processus parocciptales protruding in caudal direction; quadratum with processus orbitalis reduced; eggs incubated beneath feet; sky-pointing/wing-waving and hop displays present.
<==Sulida [Sulae, Suloidea]
|--Protoplotus [Protoplotidae]M09
| `--P. beauforti Lambrecht 1931M09
|--PhalacrocoracoideaM09
| | i. s.: Piscator Harrison & Walker 1976M09, M02
| | `--*P. tenuirostris Harrison & Walker 1976M02
| |--Borvocarbo Mourer-Chauviré et al. 2004M05
| | `--B. guilloti Mourer-Chauviré et al. 2004M09
| `--+--Anhingidae [Anhinginae, Plotinae, Plottinae, Ptynginae]PB15
| | |--Macranhinga paranensisM04
| | `--Anhinga Brisson 1760CC10 (see below for synonymy)
| | | i. s.: A. laticepsVR72
| | | ‘Plotus’ levaillantiE42
| | | A. melanogaster Pennant 1769CC10 [=Plotus melanogasterS66]
| | | |--A. m. melanogasterCC10
| | | `--A. m. vulsini Bangs 1918CC10
| | | A. nanaVR72
| | | A. pannonica (Lambrecht 1916) [=Plotus pannonicus]M02
| | | A. parvaVR72
| | | A. subvolans (Brodkorb 1956)U93
| | |--A. rufa (Dausin 1802)JT12, CC10 (see below for synonymy)
| | `--+--*A. anhingaCC10, JT12 [=Plotus anhingaCC10]
| | | |--A. a. anhingaE52
| | | `--A. a. leucogasterE52
| | `--A. novaehollandiae (Gould 1847)JT12, CC10 (see below for synonymy)
| `--+--Oligocrax stoeffelensis (Mayr 2007)PB15, M09 [=Borvocarbo stoeffelensisPB15]
| `--PhalacrocoracidaePB15
`--Sulidae [Dysporidae, Enkurosulidae, Pseudosulidae, Sulariidae, Sulinae]PB15
| i. s.: Enkurosula Kašin 1977 [=Pseudosula Harrison 1975 non Boetticher 1955]M02
| `--*E. pygmaea (Milne-Edwards 1874) [=Sula pygmaea, Microsula pygmaea, *Pseudosula pygmaea]M02
| Eostega Lambrecht 1929M02
| `--*E. lebedinskyi Lambrecht 1929M02, M09
| Empheresula Harrison 1975 [=Parasula Harrison 1975 non Mathews 1913]M02
| `--*E. arvernensis (Milne-Edwards 1867) [=Sula arvernensis, *Parasula arvernensis]M02
| Sarmatosula Grigorescu & Kessler 1977M02
| `--*S. dobrogensis Grigorescu & Kessler 1977M02
| Prophalacrocorax Harrison 1975M02
| `--*P. ronzoni (Gervais 1849)M02, M09 [=Mergus ronzoniM09, Sula ronzoniM09]
|--Masillastega Mayr 2002M05
| `--M. rectirostris Mayr 2002M09
`--+--+--Papasula abbotti (Ridgway 1893)JT12, JG19
| `--Morus Vieillot 1816JT12, CC10 [=Moris Forster 1817CC10, Sulita Mathews 1915CC10]
| | i. s.: M. olsoni Grigorescu & Kessler 1988M02
| |--*M. bassanus (Linnaeus 1758)CC10, JT12, CS77 [=Pelecanus bassanusCC10, Sula bassanaCS77, *Sulita bassanaCC10]
| `--+--M. capensis (Lichtenstein 1823)JT12, CC10 [=Dysporus capensisCC10, Sula capensisCC10]
| `--M. serrator (Gray in Dieffenbach 1843)JT12, CC10 (see below for synonymy)
`--Sula Brisson 1760JT12, CC10 [=Dysporus Illiger 1811CC10; incl. Hemisula Mathews 1913CC10, Parasula Mathews 1913CC10]
| i. s.: S. abbottiCS77
| S. cyanops (Sundevall 1837) [=Dysporus cyanops]CC10
| S. fiber Gray 1862 (n. d.)CC10
| S. fusca Vieillot 1825CC10
| S. guanoFP64
| S. piscator (Linnaeus 1758) (n. d.) [=Pelecanus piscator]CC10
|--S. sula (Linnaeus 1766)JT12, CC10 [=Pelecanus sulaCC10]
| |--S. s. sulaI92
| `--S. s. rubripes Gould 1838I92
`--+--*S. leucogaster (Boddaert 1783)CC10, JT12, CC10 [=Pelecanus leucogasterCC10]
| |--S. l. leucogasterCC10
| |--S. l. brewsteri Goss 1888CC10
| |--S. l. etesiaca Thayer & Bangs 1905CC10
| `--S. l. plotus (Forster in Lichtenstein 1844) (see below for synonymy)CC10
`--+--+--S. dactylatra Lesson 1831JT12, CC10
| | |--S. d. dactylatraCC10
| | |--S. d. bedouti Mathews 1913 [=*Parasula bedouti]CC10
| | |--S. d. melanops Heuglin 1859CC10
| | |--S. d. personata Gould 1846 [=Parasula dactylatra perstonata]CC10
| | `--S. d. tasmani van Tets, Meredith et al. 1988 (see below for synonymy)CC10
| `--S. granti Rothschild 1902JT12, CC10
`--+--S. nebouxiiJT12
`--S. variegataJT12
Nomen nudum: Sula rozieri Lucazeau 1959M02
Anhinga Brisson 1760CC10 [incl. Plotus Linné 1766B94, Plottus (l. c.)B94, Ptynx Möhring 1752 (pre-Linnean) non Blyth 1840B94]
Anhinga novaehollandiae (Gould 1847)JT12, CC10 [=Plotus novaehollandiaeCC10, A. melanogaster novaehollandiaeCC10, A. rufa novaehollandiaeCC10; incl. P. novaehollandiae derbyi Mathews 1912WS48]
Anhinga rufa (Dausin 1802)JT12, CC10 [=Plotus rufusWS48, A. melanogaster rufaCC10; incl. A. melanogaster chantrei (Oustalet 1882)CS77]
Morus serrator (Gray in Dieffenbach 1843)JT12, CC10 [=Pelecanus serratorCC10, Dysporus serratorCC10, Moris serratorCC10, Sula serrataCC10, Sula bassana serratorCC10, Sulita serratorCC10; incl. Sula australis Gould 1841 non Stephens 1826CC10, Jula (l. c.) australisCC10, Sula serrator dyotti Mathews 1913CC10, Sulita serrator rex Mathews & Iredale 1921CC10, Morus serrator rexCC10, Sula serrator rexCC10, Pelecanus sectator Solander in Medway 1993CC10]
Sula dactylatra tasmani van Tets, Meredith et al. 1988 [incl. S. dactylatra fullagari O’Brien & Davies 1990]CC10
Sula leucogaster plotus (Forster in Lichtenstein 1844) [=Pelecanus plotus, Hemisula leucgaster plotus; incl. S. leucogaster rogersi Mathews 1913, *H. rogersi]CC10
*Type species of generic name indicated
References
[B94] Bock, W. J. 1994. History and nomenclature of avian family-group names. Bulletin of the American Museum of Natural History 222: 1–281.
[CC10] Checklist Committee (OSNZ). 2010. Checklist of the Birds of New Zealand, Norfolk and Macquarie Islands, and the Ross Dependency, Antarctica 4th ed. Ornithological Society of New Zealand and Te Papa Press: Wellington.
[CS77] Cramp, S., & K. E. L. Simmons (eds) 1977. Handbook of the Birds of Europe, the Middle East and North Africa: The Birds of the Western Palaearctic vol. 1. Ostrich to Ducks. Oxford University Press: Oxford.
[E52] Eisenmann, E. 1952. Annotated list of birds of Barro Colorado Island, Panama Canal Zone. Smithsonian Miscellaneous Collections 117 (5): 1–62.
[E42] Ewer, W. 1842. List of a collection of birds from India. Proceedings of the Zoological Society of London 10: 91–93.
[FP64] Fisher, J., & R. T. Peterson. 1964. The World of Birds: A comprehensive guide to general ornithology. Macdonald: London.
Friesen, V. L., & D. J. Anderson. 1997. Phylogeny and evolution of the Sulidae (Aves: Pelecaniformes): a test of alternative modes of speciation. Molecular Phylogenetics and Evolution 7 (2): 252–260.
[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.
[JG19] James, D. J., P. T. Green, W. F. Humphreys & J. C. Z. Woinarski. 2019. Endemic species of Christmas Island, Indian Ocean. Records of the Western Australian Museum 34 (2): 55–114.
[JT12] Jetz, W., G. H. Thomas, J. B. Joy, K. Hartmann & A. Ø. Mooers. 2012. The global diversity of birds in space and time. Nature 491: 444–448.
Mayr, G. 2002. A skull of a new pelecaniform bird from the Middle Eocene of Messel, Germany. Acta Palaeontologica Polonica 47 (3): 507–512.
[M04] Mayr, G. 2004. Tertiary plotopterids (Aves, Plotopteridae) and a novel hypothesis on the phylogenetic relationships of penguins (Spheniscidae). Journal of Zoological Systematics and Evolutionary Research 43: 61–71.
[M05] Mayr, G. 2005. The Paleogene fossil record of birds in Europe. Biological Reviews 80: 515–542.
[M09] Mayr, G. 2009. Paleogene Fossil Birds. Springer.
[M02] Mlíkovský, J. 2002. Cenozoic Birds of the World. Part 1: Europe. Ninox Press: Praha.
Mlíkovský, J. 2007. Taxonomic identity of Eostega lebedinskyi Lambrecht, 1929 (Aves) from the middle Eocene of Romania. Annalen des Naturhistorischen Museums in Wien 109A: 19–27.
Nelson, J. B. 2006. Pelicans, Cormorants, and Their Relatives: The Pelecaniformes. Oxford University Press.
Olson, S. L. 1984. A brief synopsis of the fossil birds from the Pamunkey River and other Tertiary marine deposits in Virginia. In: Ward, L. W., & K. Krafft (eds) Stratigraphy and Paleontology of the Outcropping Tertiary Beds in the Pamunkey River Region, Central Virginia Coastal Plain: Guidebook for the Atlantic Coastal Plain Geological Association 1984 field trip pp. 217–223. Atlantic Coast Plain Geological Association.
Pitman, R. L., & J. R. Jehl, Jr. 1998. Geographic variation and reassessment of species limits in the “masked” boobies of the eastern Pacific Ocean. Wilson Bulletin 110: 155–170.
[PB15] Prum, R. O., J. S. Berv, A. Dornburg, D. J. Field, J. P. Townsend, E. M. Lemmon & A. R. Lemmon. 2015. A comprehensive phylogeny of birds (Aves) using targeted next-generation DNA sequencing. Nature 526: 569–573.
[S66] Schlegel, H. 1866. Communication from, on mammals and birds collected in Madagascar. Proceedings of the Zoological Society of London 1866: 419–426.
Stucchi, M., & M. Urbina. 2004. Ramphastosula (Aves, Sulidae): a new genus from the early Pliocene of the Pisco. Journal of Vertebrate Paleontology 24 (4): 974–978.
[U93] Unwin, D. M. 1993. Aves. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 717–737. Chapman & Hall: London.
[VR72] Vickers Rich, P. 1972. A fossil avifauna from the Upper Miocene Beglia Formation of Tunisia. Notes du Service Géologique 35: 29–66.
[WS48] Whittell, H. M., & D. L. Serventy. 1948. A systematic list of the birds of Western Australia. Public Library, Museum and Art Gallery of Western Australia, Special Publication 1: 1–126.