Lagenorhynchus albirostris J. E. Gray 1846
Lagenorhynchus albirostris J. E. Gray, 1846a White-beaked Dolphin Delphinus Tursio Brightwell, 1846:21, plate 1. Not Delphinus tursio Fabricius, 1780. Lagenorhynchus albirostris : J. E. Gray. 1846a:84. No type locality specified, but stated as “North Sea, coast of Norfolk ” by Gray (1846b:35); restr...
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| Summary: | Lagenorhynchus albirostris J. E. Gray, 1846a White-beaked Dolphin Delphinus Tursio Brightwell, 1846:21, plate 1. Not Delphinus tursio Fabricius, 1780. Lagenorhynchus albirostris : J. E. Gray. 1846a:84. No type locality specified, but stated as “North Sea, coast of Norfolk ” by Gray (1846b:35); restricted by Gray (1850) to Great Yarmouth. First use of current name combination. Delphinus albirostris : J. E. Gray, 1846b:35, plate 10. Incorrect reference to Lagenorhynchus albirostris (Gray, 1846a) Delphinus pseudotursio Reichenbach, 1846:plate 24, figure 76. New name based on Brightwell’s (1846) description (see “Nomenclatural Notes”). Delphinus Ibsenii Eschricht, 1846:297. Type locality: “Agger Tange,” West coast of Jutland, Denmark. Delphinus (Lagenorhynchus) albirostris: van Beneden, 1860:28. Name combination. CONTEXT AND CONTENT. Order Cetacea, suborder Odontoceti, family Delphinidae. Subfamily assignment remains unsettled. Traditionally, the genus Lagenorhynchus comprised 6 shortsnouted delphinid species. Molecular research, however, has found this grouping to be polyphyletic, that is, comprising at least 2 unrelated lineages: (1) the Pacific and Southern ocean members ( L. obliquidens , and L. obscurus , L. australis , and L. cruciger , respectively), assigned to the subfamily Lissodelphininae; and (2) the 2 North Atlantic members of the genus ( L. albirostris and L./ Leucopleurus acutus — Cipriano 1997; LeDuc et al. 1999; Harlin-Cognato and Honeycutt 2006; McGowen et al. 2009; McGowen 2011; Banguera-Hinestroza et al. 2014). Molecular studies addressing the relationship between the latter 2 species have provided conflicting results (LeDuc et al. 1999; Harlin-Cognato and Honeycutt 2006; Banguera-Hinestroza et al. 2014). All studies agree that the taxa have basal positions in the Delphinidae phylogeny, but they either represent separate lineages (subfamilies) within the dolphin family (Delphinidae) or they are indeed sister species and form their own subfamily Lagenorhynchinae (Banguera- Hinestroza et al. 2014). In the latter case, the time of their divergence has been estimated as far back as 11.49 million years ago, which may merit assignment to separate genera (Banguera-Hinestroza et al. 2014). NOMENCLATURAL NOTES. Brightwell’s (1846) description of a dolphin from Great Yarmouth that he assigned to Delphinus Tursio of Fabricius (1780) was promptly corrected by Gray (1846a) in what is considered the formal description of the white-beaked dolphin. In this and his later paper (1846b), Gray applies Lagenorhynchus as the generic name, although in his 2nd account (Gray 1846b), he incorrectly credited himself for applying the name Delphinus albirostris in the 1st account (Gray 1846a). Later, the binomen Delphinus albirostris was used by Gray and many others. The work of Reichenbach (1846) does not include a description of the new species in the text but contains on plate XXIV, as number 76, a mirrored version of Brightwell’s (1846) original figure with a caption reading: Delphinus pseudotursio Rchb. D. Tursio Th. Brightwell. An. Mag Hist. 1846 . Hereby Reichenbach, independently from Gray (1846a, 1846b) recognized Brightwell’s dolphin as a new species. The new species name chosen by Reichenbach, clearly is based on Brightwell’s specimen and therefore available, but since no exact date of publication could be established it falls into junior synonymy. Wagner (1846) provided a short Latin description of the species and in the accompanying plate volume (Wagner 1847), a mirrored version of the original Brightwell (1846) drawing. He considered Lagenorhynchus a subgeneric name introduced by Gray and consequently named it D [ elphinus ]. albirostris. Generic and subgeneric names were often used interchangeably. van Beneden (1860) for the 1st time gave the scientific name in full: Delphinus ( Lagenorhynchus ) albirostris , but later, he used both Delphinus albirostris and Lagenorhynchus albirostris . Eschricht (1846) gave a short description of the skeleton of a new dolphin species he intended to describe under the name Delphinus Ibsenii , but he never published an extended description. Gray (1850) and Claudius (1853) put Delphinus Ibsenii into the synonymy of Lagenorhynchus albirostris . Kinze (2009) established that Gray’s specific name ( albirostris ) antedates Eschricht’s and therefore is to be considered the senior synonym. Delphinus tursio was still in use for specimens of whitebeaked dolphins until at least 1861 (Sundevall 1861). Several European museums hold specimens of the white-beaked dolphin originating from waters around Greenland and collected before the formal description in 1846, these were originally assigned to Delphinus delphis. The name, Lagenorhynchus , is composed of the Latin word lagena , meaning bottle or flask, and the Greek word rhynchus , meaning snout. The specific name, albirostris (Latin: whitebeaked), refers to the common, but not universal, white beak. Additional common names for L. albirostris include the following: hvidnaese (Danish); witsnuitdolfijn (Dutch); kjarthvitur springari (Faroese); dauphin avec bec blanc (French); Weissschnauzendelfin (German); ardluarsuk (Greenlandic); Blettahnýðir or Hnýðingur (Icelandic); deilf na ngoba bána (Irish); kvitnos (Norwegian); delfin bialosy (Polish); belomordyi del’fin (Russian); delfin de hocicio blanco (Spanish); vitnosdelfin (Swedish). It is commonly referred to as “springer” or “jumper,” especially in parts of Canada, the Faroes, and Norway. These latter names are also used to refer to the Atlantic whitesided dolphin. DIAGNOSIS Lagenorhynchus albirostris is most likely to be confused with the Atlantic white-sided dolphin, L . acutus , from which it differs by the white or gray beak, a light gray-to-white saddle behind the dorsal fin, and the lack of a yellow field on the posterior flank. Skulls of L . albirostris can be distinguished from those of other North Atlantic delphinids by a toothrow with more than 20 teeth and by the shorter (<240 mm) and wider (> 125 mm) rostrum. Skeletons of L . albirostris differ from those of other cetaceans by the high vertebral count, 88–93. Only right whale dolphins ( Lissodelphis —up to 90 vertebrae) and Dall’s porpoise ( Phocoenoides dalli —up to 98 vertebrae), both absent in the North Atlantic, reach this range. GENERAL CHARACTERS Lagenorhynchus albirostris is robust compared to most other delphinids, with a comparatively short, thick beak protruding 5–8 cm from the offset of the melon. The tall, falcate dorsal fin is located around mid-length. Coloration is complex and changes during ontogeny, this has been described and broken into components by Bertulli et al. (2016). As the vernacular name implies, white-beaked dolphins have a white or whitish beak, which in many, particularly adult individuals, is darkened to an ashy gray (Bertulli et al. 2016). The color of the beak often continues backward a few cm posterior to the base of the melon. The dorsum, flanks, and the remainder of the melon are dark gray, whereas the dorsal fin, flippers, and fluke are generally dark gray except for a white or light gray saddle behind the dorsal fin and the occurrence of light patches on the anterior leading edges of fin and flippers in adults. A light patch surrounds the eye, with a darker section immediately around the eye. The dark gray areas of the dorsum and flanks are overlaid by a lighter gray blaze starting near the anal region and extending dorso-anteriorly along the peduncle and flank to the level of the flippers. Another light gray area on the anterior (thoracic) flank lies between the flipper and the flank patch. The belly is entirely light gray-to-white with a narrow white medial band from the flippers to the genital slit (Figs. 1 and 2). This general pattern varies among ontogenetic stages, individuals, and possibly geographic areas (Harmer 1927; Mercer 1973; Camphuysen 1991; Ree 1994; Bertulli et al. 2016). The longest known specimen is a 310 cm male from the British Isles (Fraser 1974), but few specimens reach 300 cm in total length (Guldberg and Nansen 1894; Harmer 1927; Galatius et al. 2013a). Adult males are larger than females (Dong et al. 1996) due to an extended period of growth (Galatius et al. 2013a). In a sample from the North Sea, adult females were 242–265 cm with a mean length of 251 cm and males were 252–290 cm with a mean of 273 cm (Galatius et al. 2013a). Ranges of selected body measurements (cm; by C. C. Kinze) from adult females and males, respectively, stranded in the eastern North Sea were: dorsal fin height, 22–38 ( n = 19) and 26–40 ( n = 15); anterior flipper length, 32–48 ( n = 21) and 36–52 ( n = 19); fluke span, 42–70 ( n = 18) and 52–83 ( n = 17); and snout to center of genital aperture, 90–120 ( n = 15) and 75–90 ( n = 16). The observation of a pregnant female only 174 cm long suggests that L . albirostris may reach adulthood at smaller sizes in the western North Atlantic (Dong et al. 1996). Few weights have been reported, the largest known male weighed 354 kg (van Utrecht 1981) and the largest female 306 kg (Reeves et al. 1999). Means and ranges (mm; by A. Galatius) of selected cranial measurements of adult females and males, respectively, from the eastern North Sea were: condylobasal length, 444 (415–452; n = 16), 451 (435–464; n = 12); rostral length, 219 (206–227; n = 17), 220 (212–226; n = 12); rostral width at base, 139 (130–145; n = 16), 143 (132–153; n = 12); width at antorbital processes, 228 (216–241; n = 16), 234 (222–246; n = 12); zygomatic width, 243 (233–257; n = 16), 253 (240–266; n = 12); right mandible length, 369 (350–385; n = 17), 376 (354–391; n = 15); length of right upper toothrow, 192 (182–202; n = 16), 194 (188–202; n = 12). The skull has large, deep temporal fossae and robust antorbital, postorbital, and zygomatic processes (Fig. 3). DISTRIBUTION Lagenorhynchus albirostris is endemic to the cooler temperate and subarctic North Atlantic (Reeves et al. 1999; Fig. 4). Four principal centers with higher density have been identified (Kinze 2009): (1) The Labrador Shelf and southwestern Greenland; (2) around Iceland; (3) around the northern part of the British Isles and the North Sea; (4) the shelf along the Norwegian coast, including the Barents Sea. In (1), the species has been recorded from western Greenland in the North (71°) to Cape Cod, Massachusetts (Alling and Whitehead 1987; Kingsley and Reeves 1998; Waring et al. 2008; Lawson and Gosselin 2009; Hansen and Heide-Jørgensen 2013). Apart from 2 recent sightings near the southeastern coast of Baffin Island, the species is unknown in this area (Reinhart et al. 2014). In (2), L . albirostris is the most common dolphin (Vikingsson and Ólafsdóttir 2004; Pike et al. 2009). This area may also extend to southeastern Greenland, where it is also the most common dolphin (Jonsgård and Christensen 1968). In (3), the highest sighting rates are from the Outer Hebrides and in the western North Sea, but its distribution also extends into the central North Sea to the Danish coast, around Ireland and even some sightings in the English Channel (Northridge et al. 1995, 1997; Hammond et al. 2002; Reid et al. 2003; Hammond et al. 2013). In (4), distribution of L . albirostris extends throughout the western Barents Sea except for the northernmost parts (Øien 1996; Fall and Skern-Mauritzen 2014). FOSSIL RECORD There is no fossil record of the taxon. Subfossil remains from the greater North Sea and Baltic Sea area document its presence in European waters since the last glacial period (Aaris-Sørensen et al. 2010). Degerbøl (1936) reported subfossils from the medieval Norse West Settlement in Greenland. FORM AND FUNCTION The dental formula of Lagenorhynchus albirostris is 25–28 homodont teeth in each jaw, but the 3 anteriormost teeth often do not protrude from the gums (van Bree and Nijssen 1964). Vertebral formula is 7 C, 15–16 T, 23–24 L, 43–45 Ca, total 88–92 vertebrae (Lütken 1888; True 1889) or 7 C, 14 T, 27 L, 45 Ca, total 93 (Tomilin 1967). This is the highest count of vertebrae in a delphinid. The vertebrae have short, wide centra and long spinal and transverse processes. Such vertebral morphology is an adaptation to fast, dynamic swimming at the expense of flexibility (Buchholtz 2001). We have observed that the first 2 cervical vertebrae are always fused, but often all 7 are fused. The phalangeal formula is I2, II6, III4, IV1, V0 (Tomilin 1967). In most specimens, the right humerus is longer and more robust than the left, indicating lateralized behavior, something that has been recorded in several cetacean species (Galatius 2006). The 2–4 vibrissae on each upper lip in fetuses and neonates (Turner 1889; Japha 1912) disappear during development as in other Odontoceti and adults have none. The integumentary blubber is 18–31 mm thick, except for the fins and flukes in which a countercurrent heat exchange system operates to conserve heat (Scholander and Schevill 1955). The organ systems of L . albirostris exhibit a typical delphinid organization (muscular system—Murie (1873); nervous system: spinal cord—Cunningham (1877), brain—Weber (1887); sensory system—Fraser (1952); lungs—Cleland (1884) and Weber (1887); digestive system: stomach—Clark (1876), Cleland (1884), Weber (1887), and Turner (1889), small and large intestine—van Beneden (1860) and Cleland (1884); circulatory system: blood vessels—Slijper (1936), heart—Smet et al. (1985) and van Nie (1985, 1986); female reproductive system—van Beneden (1860) and Meek (1918); functional anatomy of mammary slits—Gylling (1912)). In animals from the North Sea ( n = 10), the lungs weighed 1,100 –2,000 g (left) and 1,300 –2,200 g (right). Heart masses were 800–1,900 g, liver masses 1,800 –3,400 g, and kidneys weighed 300–600 g each. Adrenals are tiny: 2–8 g (Reeves et al. 1999; and measured by C. C. Kinze). ONTOGENY AND REPRODUCTION Ontogeny. —Some highly specialized information is known only from single fetal specimens of Lagenorhynchus albirostris (larynx—Thompson 1890; retina—ten Doesschate 1918). Length at birth is estimated at 110–120 cm (Tomilin 1967; Collet and Duguy 1981), but the age and length at weaning are not known. Growth has only been thoroughly investigated in a North Sea sample of 64 females and 52 males (Galatius et al. 2013a). During the first 3 years, young of both sexes grew to about 210 cm, after which growth rate slowed somewhat. Females reached 95% of asymptotic length by about 5.2 years but males continued growing 5 more years, resulting in sexual size dimorphism. Females become sexually mature at 6–10 years and 230–240 cm in length. Males become sexually mature about 2 years later than females and are typically 230–260 cm in length. Adult size of the cranium is achieved at about 6 years of age in the North Sea population ( n = 81), whereas sutures of the skull are not fully developed until about 9 years of age (Galatius 2010). Reproduction. —Paired testes mass in 16 adults from the North Sea showed annual fluctuations from about 700 g during the winter to near 2,100 g during the boreal summer, suggesting that mating occurs in the latter period (Galatius et al. 2013a). Around the United Kingdom, strandings of calves below 130 cm in length are reported almost exclusively between June and September, supporting that parturition occurs during this time (Fraser 1974; Canning et al. 2008). Gestation is believed to last about 11 months (Reeves et al. 1999), litter size is 1 as in other Cetacea. Only 3 of 22 sexually mature females were lactating and only 1 was pregnant in the North Sea sample of Galatius et al. (2013a). This observation indicates either a significant resting period between parturitions or segregation of reproductively active females from others. ECOLOGY Population characteristics. —Skull morphometric evidence suggesting segregation between the eastern and western North Atlantic populations of Lagenorhynchus albirostris (Mikkelsen and Lund 1994) was confirmed by a molecular study (Banguera-Hinestroza et al. 2010), which also found segregation between dolphins from the United Kingdom and North Sea and those from northern Norway. The latter study did not include samples from Iceland or Greenland; thus, population structure throughout the range of the species remains unresolved. Using> 50,000 single nucleotide polymorphisms, Fernandez et al. (2016) detected 2 genetic clusters among 70 L. albirostris sampled throughout the eastern North Atlantic. Aerial surveys conducted from 1978 to 1982 for the shelf waters from Cape Hatteras to Nova Scotia yielded a population abundance estimate of 573 ( CV : 0.69— Cetacean and Turtle Assessment Program 1982). Aerial surveys in the 1980s off eastern Newfoundland and the Labrador coast yielded an estimate of 5,500 L . albirostris (no CV —Alling and Whitehead 1987). An abundance of 3,486 individuals (95% CI : 2,001 –4,971) was estimated from a shipboard survey of a small segment of the Labrador shelf in 1982 (Alling and Whitehead 1987). Aerial surveys in 1995 and 1996 in the Gulf of St. Lawrence yielded estimates of 2,640 ( SE : 2,080) and 2,380 ( SE : 2,540), respectively (Kingsley and Reeves 1998), and a 2007 aerial survey of the Canadian shelf from north Labrador to the United States border gave a preliminary estimate of 1842 ( CV : 0.22—Lawson and Gosselin 2009). A coast-to- 90 km aerial survey of the shelf of West Greenland up to 71°N yielded an estimate of 11,984 ( CV : 0.19) L . albirostris in 2007 (Hansen and Heide-Jørgensen 2013). Aerial surveys in Icelandic waters in 2001 yielded a point estimate of 31,653 (95% CI : 17,679 –56,672 — Pike et al. 2009). This estimate also included other dolphin species, although the overwhelming majority of the sightings were L . albirostris . In the greater North Sea, shipboard surveys in 1994 yielded an estimate of 7,856 individuals ( CV : 0.30— Hammond et al. 2002). A 2005 estimate for this area was 16,536 ( CV : 0.30— Hammond et al. 2013). The latter survey covered an area to the west that was not covered in 1994 and showed high density in 2005. The design of this survey was criticized for being inappropriate for the patchy distribution of L . albirostris (MacLeod 2014). In their response, Hammond et al. (2014) maintained that the design was appropriate for large-scale abundance estimates, but not f : Published as part of Galatius, Anders & Kinze, Carl Christian, 2016, Lagenorhynchus albirostris (Cetacea: Delphinidae), pp. 35-47 in Mammalian Species 48 (933) on pages 35-43, DOI: 10.1093/mspecies/sew003, http://zenodo.org/record/4573390 : {"references": ["GRAY, J. E. 1846 a. On the British Cetacea. Annals and Magazine of Natural History 17: 82 - 85.", "BRIGHTWELL, T. 1846. Observations on a specimen of the bottlenosed dolphin, Delphinus Tursio, Fabr., taken at Great Yarmouth, October 1845. The Annals and Magazine of Natural History 17: 21 - 22.", "FABRICIUS, O. 1780. Fauna Groenlandica: systematice sistens animalia Groenlandiae occidentalis hactenus indagata, quad nomen specificum, triviale, vernaculumque: synonyma auctorum plurium, descriptionem, locum, victum, generationem, mores Othonis Fabricii. Ioannis Gottlob Rothe, Copenhagen, Denmark.", "GRAY, J. E. 1846 b. 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