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A New Species of Shearwater of the Genus Calonectris (Aves: Procellariidae) from a Middle Pleistocene Deposit on Bermuda

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A New Species of Shearwater of the Genus Calonectris (Aves: Procellariidae) from a Middle Pleistocene Deposit on Bermuda PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 121(3):398^09. 2008. A new species of shearwater of the genus Calonectris (Aves: Procellariidae) from a middle Pleistocene deposit on Bermuda Storrs L. Olson Division of Birds, Department of Vertebrate Zoology, National Museum of Natural History, P.O. Box 37012, Smithsonian Institution, Washington, D.C. 20013-7012, U.S.A., e-mail: [email protected] Abstract.•Remains of at least 26 individuals of a Calonectris shearwater were recovered from a Pleistocene beach deposit on Bermuda that formed when sea-level was more than 21 m above present level during an interglacial (Marine Isotope Stage 11) 400,000 yr ago. Two prefledging juveniles in the sample indicate breeding on the island. This shearwater was the general size of C. d. diomedea of the Mediterranean but differs in proportions and in quahtative characters and is described as Calonectris wingatei, new species. The species appears to have become extinct shortly after the time of deposition, when rising sea-level is also thought to have caused the extinction of the Short-tailed Albatross Phoebastria albatrus on Bermuda. The Neogene history of the taxa of Calonectris in the Atlantic basin is examined in the context of major geological and océanographie events. Bermuda's isolated position in the water Puffinus puffinus, but this species North Atlantic, 1000 km east of the was apparently never an established North American mainland, places it resident, as the abundant fossil record within foraging distance for procellarii- from the island (Olson et al. 2005) has so form birds of both temperate and sub- far yielded only a single sternum of the tropical waters as well as hydrographie species (Olson 2004a). Finally, the Short- fronts that form along the nearby Gulf tailed Albatross Phoebastria albatrus once Stream (Haney & McGillivary 1985). bred on Bermuda, but the Atlantic Despite this, the diversity of breeding population appears to have been extir- Procellariiformes known from the island pated by the maximum rise of sea level is quite low. The highly endangered (>+21 m) in the Pleistocene 400,000 yr Cahow Pterodroma cahow, once the most ago during Marine Isotope Stage (MIS) abundant bird on Bermuda, is the only 11 (Olson & Hearty 2003). The former member of the order that persists on the presence of a fifth species of procellarii- island. A small extinct shearwater, which form on Bermuda is here documented provisionally takes the name Puffinus from a large assortment of fossil bones of parvus Shufeldt (see Olson 2004a), is a shearwater of the genus Calonectris known only from fossils, and appears to from a single deposit that formed at the have been succeeded, probably in historic height of the same sea level rise that is times, by the West Indian Audubon's postulated to have wiped out the alba- Shearwater P. Iherminieri, which in turn tross. was extirpated on Bermuda in the 20th Three living taxa of shearwaters of the century (Amos 1991). There are three genus Calonectris currently breed on known nesting attempts by Manx Shear- islands in the North Atlantic and Medi- VOLUME 121, NUMBER 3 399 terranean, all of which had at one time were made with: Calonectris d. borealis• been considered subspecies of a single Massachusetts 17767, 17770, 17773-76, species. Of these, it is now generally 17788-91, 17831, New Jersey 347825; C. agreed that the Cape Verde Shearwater d. diomedea•Greece 488776-77, 555936 C. edwardsi is a distinct species by virtue (several individuals), Italy 612133, North of much smaller size and plumage differ- Carohna 620710-18; C. edwardsi (trunk ences (Hazevoet 1995:67), as well as by skeletons only)•Razo I., Cape Verdes molecular differences (Gomes-Diaz et al. 560762-68; and C. leucomelas•Halma- 2006). The Mediterranean or Scopoli's hera, Indonesia 558350. Fossil material is Shearwater C. diomedea breeds on islands housed in the Department of Paleobiolo- in the Mediterranean Sea. The larger gy, Smithsonian Institution. Measure- Cory's Shearwater C. borealis breeds in ments were taken with digital calipers to the North Atlantic islands of the Azores, the nearest 0.1 mm. Canaries, Salvages, Madeira, and the Berlengas Islands off of Portugal. There Family Procellariidae has been much variation in opinion as to whether this taxon deserves full specific Genus Calonectris Mathews & Iredale, rank or should be regarded as a subspe- 1915 cies of C. diomedea (discussed beyond). I originally interpreted the Pleistocene The species of Calonectris were usually accumulation from Bermuda as having placed in the genus Puffinus, even after been derived from beached wintering their separation had been proposed (e.g., individuals of C. d. diomedea (Olson & Peters 1931). It was only following the Rasmussen 2001, Olson et al. 2005). With innovative study of Kuroda (1954) that further study, I have found that these the validity of Calonectris came to be fossils belong to a previously unrecog- accepted. The long, slender rostrum allies nized species, similar in size to C. d. the Bermuda species with the Puffininae diomedea but with larger hindlimbs and as opposed to the fulmarines. As noted by more robust long bones. This species was Kuroda, only in Calonectris are the breeding on Bermuda up until the middle anterior intervertebral foramina of the Pleistocene, after which it appears to have sacrum closed by ossification. In Calonec- become extinct. tris the shaft of the humérus is terete, not fiattened as in subgenus Puffinus and to a Materials and Methods lesser extent in Ardenna, but is relatively longer and more slender than in any of Modern comparative skeletal material the taxa, except Thyellodroma. The bra- was examined in the collections of the chial depression is more restricted than in Division of Birds, National Museum of Fulmarus, Pterodroma, or Procellaria, but Natural History, Smithsonian Institution, is not as small and decidedly rounded in Washington, D.C. (USNM). At the ge- shape as in Thyellodroma. In Calonectris neric level, comparisons were made with the shaft of the femur in medial view is Fulmarus glacialis 561911, Pterodroma slightly curved (straight in Fulmarus, leucoptera 615080, Procellaria aequinoc- Pterodroma, Procellaria), but decidedly tialis 490787, Calonectris d. diomedea less curved than in any of the species of 620711, Puffinus (Hemipuffinus) crea- Puffinus, except the subgenus Hemipuffi- topus 559553, P. (Thyellodroma) pacificus nus, from which Calonectris differs in 500712, P. (Ardenna) gravis 561135, P. having the shaft relatively more slender, (Puffinus) griseus 599153, P. {P.) puffinus with the appearance of being slightly 491958. At the specific level, comparisons constricted near the midpoint. The inner 400 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Cranial elements of Calonectris. A, skull of C. wingatei holotype (USNM 535555); B, mandible of C. wingatei (USNM 535566); C, D, skiill and mandible of C. d. diomedea (USNM 620716). Mandibles appear as mirror images. Arrows indicate the larger fonticulus orbitocranialis of the cranium and in the mandible the depression for the M. adductor mandibulae. cnemial crest of the tibiotarsus is much ratojugals and all elements of the palate; less developed in Calonectris than in collected 9 Aug 1984 by S. L. Olson, D. B. Puffinus. In all Puffinus, except subgenus Wingate, and Frederick V. Grady. Hemipuffinus, the shaft of the tarsome- Type locality.•Bermuda, Hamilton Pa- tarsus is more laterally compressed and rish, Calonectris Quarry, 32°20'20.9"N, the inner trochlea more retracted than in 64°42'24.5W" (see Olson et al. 2005). The Calonectris, in which the tarsometarsus is site where the fossils were collected, called much more gracile than in Hemipuffinus. Calonectris Quarry, was exposed in the In all of these respects the Bermuda Government Quarry on the western side of material is clearly referable to Calonectris. Castle Harbour on the eastern end of Bermuda. It was described briefly by Calonectris wingatei, new species Olson & Hearty (2003) and Olson et al. Figs. 1-4 (2005) and in detail by Olson & Hearty Holotype.•USNM 535555 articulated (2009) because of its importance in pro- cranium and rostrum lacking the quad- viding multiple lines of evidence for sea- VOLUME 121, NUMBER 3 401 Fig. 2. Wing elements of Calonectris. A, B, E, G, C d. diomedea. C, D, F, H, C wingatei. A-D, right humeri (USNM 620714, 620710, 535601, 535600). E, F, right ulnae (USNM 620714, 535604); G, H, left carpometacarpi (USNM 620716, 535605). Note the stouter shafts in C wingatei. level having exceeding 20 m above present localities in the Bahamas (Hearty et al. in the Middle Pleistocene. The associated 1999) and was later confirmed at another fauna (Olson & Hearty 2009) consisted of site on Oahu, Hawaiian Islands (Hearty terrestrial and marine invertebrates, fish, 2002) and elsewhere. This event would three species of seabirds in addition to have had a major impact on seabirds Calonectris (Pterodroma cahow, Puffinus worldwide. parvus, Phoebastria albatrus), three species Measurements of holotype (mm).•To- of land birds, a skink (Olson et al. 2006), tal length 101.8, length of rostrum from and a vagrant bat (Grady & Olson 2006). nasofrontal hinge 55.5, length of cranium The accumulation clearly formed as a from nasofrontal hinge 47.5, width of beach deposit, the birds having been nasofrontal hinge 13.4, least width of washed into a crevice that was probably interorbital bridge 10.4, greatest width of near the maximum of normal wave runup cranium (at postorbital processes) 35.9, on a sloping beach. The elevation of the depth of rostrum at nasofrontal hinge 11.4. site was determined by transit survey as Paratypes.•535627 rostrum and dor- being exactly 70 ft (21.3 m) above present sal surface of cranium; USNM 535552- sea level. 54, 535556 crania; 535557, 525628-30 Age.•Middle Pleistocene, Marine Iso- rostra; 535558 anterior half of rostrum; tope Stage 11, 400,000 yr BP (Olson & 535559-65 rostral hooks; 535565-78 in- Hearty 2003, 2009).
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