Steno bredanensis (Cetacea: Delphinidae) - BioOne

MAMMALIAN SPECIES 43(886):177–189 

Steno bredanensis (Cetacea: Delphinidae) 

KRISTI L. WEST, JAMES G. MEAD, AND WHITNEY WHITE 

College of Natural and Computational Sciences, Hawaii Pacific University, Kaneohe, HI 96744, USA; kwest@hpu.edu (KLW); 

volcomcurly@hotmail.com (WW) 

Division of Mammals, Smithsonian Institution, Washington, DC 20013-7012, USA; meadj@si.edu (JGM) 

Abstract: Steno bredanensis (Cuvier in Lesson, 1828) is a small odontocete commonly called the rough-toothed dolphin. A 

slender, gray dolphin with a slightly darker cape, this species is most easily distinguished from other small delphinids by a 

gradually sloping forehead and a long rostrum. It is the only species in the genus Steno. Despite reports of sightings or 

stranded specimens from all tropical and subtropical oceans, the species is thought to typically occur in low abundance. The 

conservation status of S. bredanensis is poorly known. 

Key words: cetacean, dolphin, marine mammal, odontocete, rough-toothed dolphin 

E 26 September 2011 American Society of Mammalogists 

Synonymy completed 1 June 2011 

DOI: 10.1644/886.1 

www.mammalogy.org 

Steno Gray, 1846 

Steno Gray, 1846:43. Type species Delphinus rostratus 

Cuvier, 1833 (5 Delphinus bredanensis Cuvier in Lesson, 

1828), by monotypy. 

Glyphidelphis Gervais, 1859:301. Type species Delphinus 

rostratus Cuvier, 1833 by monotypy. 

Delphinus (Steno) perspicillatus Peters, 1877:360, Taf. 3. 

Type locality ‘‘im atlantischen Ocean, in 32u 29’ 7 S. B. 

und 2u 1’ W. L. Gr. harpunirt wurde.’’ 

Steno bredanensis: Miller and Kellogg, 1955:657–658. First 

use of current name combination. 

Delphinus chamissonis: Herskovitz, 1966:17. Not Delphinus 

chamissonis Wagner, 1846. 

CONTEXT AND CONTENT. Order Cetacea, suborder Odontoceti, 

family Delphinidae, subfamily Steninae. The genus 

Steno is monotypic. 

CONTEXT AND CONTENT. 

Context as for genus. Species is monotypic. 

Steno bredanensis (Cuvier in Lesson, 1828) 

Rough-toothed Dolphin 

Delphinus rostratus (Desmarest, 1817:160). Type locality 

‘‘Paimpol, Brittany, France.’’ Preoccupied by Delphinus 

rostratus Shaw, 1801. 

Delphinus frontatus (Cuvier, 1823:278). Type locality unknown. 

Delphinus bredanensis Lesson, 1828:206. Replacement name 

for Delphinus rostratus Desmarest, 1817; preoccupied by 

Delphinus rostratus Shaw, 1801. 

Delphinus planiceps Schlegel, 1841:27, tab IV. A replacement 

name for Delphinus bredanensis van Breda, 1829. 

Delphinorhynchus santonicus Lesson, 1836:330. Type locality 

‘‘lIe d’Aix, mouth of the Charente River.’’ Type based 

on a stranded individual that was not preserved. 

Delphinus compressus (Gray, 1843:105). Nomen nudum. 

Steno rostratus Gray, 1846:30, 46. Renaming of Delphinus 

rostratus Desmarest, 1817. 

Steno compressus Gray, 1846:43, pl. 27. Type locality unknown. 

Fig. 1.—Pubertal male Steno bredanensis from Dolphin Quest 

French Polynesia, photographed in 2000. Dolphin Quest French 

Polynesia (The Moorea Dolphin Center) is a captive-care facility 

that houses dolphins in French Polynesia, located in the South 

Pacific. Used with permission of the photographer Cecile Gaspar.

178 MAMMALIAN SPECIES 43(886)—Steno bredanensis 

Hershkovitz (1966:17) follows Wagner’s (1846) attribution 

to ‘‘Wiegmann, 1841 (or earlier)’’ based on Wagner’s 

labeling of his plate CCCLIX (359) as ‘‘Delphinus Chamissonis 

Wiegm.’’ We surmise that Wagner had seen a 

manuscript by Wiegmann because we were not able to 

locate the record in the published literature. 

DIAGNOSIS 

Fig. 2.—Dorsal, ventral, and lateral views of the skull and lateral 

view of mandible of an adult female Steno bredanensis (National 

Museum of Natural History 572792). Note that view of the right 

mandible has been reversed to align it with the cranium. Specimen 

is from Wreck Island, Virginia. Condylobasal length of skull is 

503 mm. Used with permission of the photographer Michael Potter. 

NOMENCLATURAL NOTES. The historical nomenclature of 

S. bredanensis is particularly confusing. Schevill (1987) and 

Flower (1884) have both provided accounts of this history. 

The fate of the type specimen is unclear and it may have 

been lost. The relationship between S. perspicillatus and S. 

bredanensis is addressed by Fraser (1966). 

A series of illustrations of the external appearance and 

skull of the specimen of Delphinus bredanensis from Brest that 

Lesson (1828:figure 1) described and an account of Cuvier’s 

role in the historical nomenclature of S. bredanensis is 

provided by van Breda (1829:235, 236, 238, plate I—figures 

1–6, plate II—figures 1 and 2). This is the paper that is 

miscited by Hershkovitz (1966:16), following Flower 

(1884:484) as the authority for D. planiceps van Breda. The 

trivial name planiceps does not occur in van Breda (1829). 

In the wild, Steno bredanensis can most easily be 

distinguished from other small delphinids by a forehead 

that gradually slopes into a long and slender rostrum, 

lacking the demarcation that separates the melon and beak 

(Fig. 1). Although Stenella and the Indo-Pacific bottlenose 

dolphin (Tursiops aduncus) have a similarly long rostrum, 

they have the clear demarcation that is absent in S. 

bredanensis. The bottlenose dolphin (T. truncatus) has a 

much shorter rostrum and a dorsal fin that is less erect but 

may still be confused with S. bredanensis (Jefferson and 

Leatherwood 1993; Perrin et al. 2007). 

The color pattern appears to vary according to 

geographic location and age of the individual (Miyazaki 

and Perrin 1994). S. bredanensis is gray in color with a 

slightly darker gray cape and often has distinctive white lips 

(Baker 1987; Jefferson and Leatherwood 1993). The darker 

cape is apparent from above the eye to the dorsal fin, 

gradually widening, and extending down the sides of the 

animal (Miyazaki and Perrin 1994; Fig. 1). The sides of the 

animal are typically a lighter gray color, and there may be 

white, pink, or yellow splotches or scarring on the tip of the 

rostrum, along the lower jaw, and in the ventral region. The 

mottled appearance is more frequently seen in older 

individuals, whereas uniform shades of gray are common 

in younger animals (Miyazaki and Perrin 1994). 

The skull (Fig. 2) may be confused with that of the 

Indo-Pacific humpback dolphin (Sousa chinensis) orStenella 

but the skull of S. bredanensis is generally longer than those 

of Stenella (condylobasal length . 472 mm—Miyazaki and 

Perrin 1994). The rostrum of S. bredanensis is also long, 

approximately 60% of the condylobasal length, similar to the 

ratio between condylobasal length and rostrum in the 

spinner dolphin (Stenella longirostris), Pantropical spotted 

dolphin (Stenella attenuata), and striped dolphin (Stenella 

coeruleoalba—Perrin and Gilpatrick 1994; Perrin and Hohn 

1994; Perrin et al. 1994). The mandibular symphysis of S. 

bredanensis is about one-third the length of the mandible, 

whereas the mandibular symphysis does not generally exceed 

30% of the mandibular length in Sousa (Ross et al. 1994). 

The orbits of S. bredanensis also are relatively large (. 13% 

of condylobasal length) compared to Sousa. There are 19–26 

teeth on each side in the upper jaw and 19–28 teeth in the 

lower jaw (Miyazaki and Perrin 1994). Tooth counts also 

can be used to distinguish between S. bredanensis and Sousa 

because S. chinensis has 30–38 teeth in each jaw and alveoli

43(886)—Steno bredanensis MAMMALIAN SPECIES 179 

Fig. 4.—Geographic distribution of Steno bredanensis. 

Fig. 3.—Magnified diagram of teeth ridges of Steno bredanensis 

reprinted from Neuville (1928), illustrating the fine tooth ridges 

that are characteristic of S. bredanensis. The tooth is from the 

middle of the upper jaw; lingual aspect on the left, medial aspect on 

the right. 

that are larger and more widely spaced (Van Waerebeek 

et al. 1999). The common name comes from fine, longitudinal 

ridges apparent on the surface of the teeth (Fig. 3). 

This character can be diagnostic when comparing teeth from 

S. bredanensis side by side with teeth from other delphinids. 

GENERAL CHARACTERS 

Steno bredanensis is sexually dimorphic, with males 

larger than females (Miyazaki and Perrin 1994; West 2002). 

The total length of 64 male and female adult specimens 

compiled from the Pacific and Atlantic oceans and the 

Mediterranean Sea ranged from 209 to 265 cm (Miyazaki 

and Perrin 1994). Larger specimens are reported from Brazil, 

where maximum length for a male is 283 cm and for a female 

270 cm (Siciliano et al. 2007). 

Average body mass for S. bredanensis is reported as 

130 kg (Watson 1981). Fourteen specimens ranged from 90 

to 155 kg (Miyazaki and Perrin 1994). The total body length 

(L)–total body weight (W) relationship for a combination of 

15 males and females is estimated as L 5 3.123 + 2.805 log 

W(r 5 0.999—Miyazaki and Perrin 1994). 

Selected external measurements of adult specimens compiled 

from the Pacific and Atlantic oceans and the Mediterranean 

Sea were as follows (cm): tip of upper jaw to apex of melon, 

10–14 (n 5 15); tip of upper jaw to end of gape, 22–37 (n 5 35); 

tip of upper jaw to umbilical scar, 79–121 (n 5 25); tip of upper 

jaw to tip of dorsal fin, 129–131 (n 5 2); girth at axilla, 100–110 

(n 5 17); anterior length of flipper, 36–49 (n 5 36); width of 

flipper, 13–17 (n 5 33); span of flukes, 46–65 (n 5 35); width of 

fluke, 14–23 (n 5 27); and height of dorsalfin,18–28(n 5 23— 

Miyazaki and Perrin 1994). 

The characteristically long rostrum of S. bredanensis is 

apparent in the skull and mandible (Fig. 2). Cranial 

measurements compiled from a number of sources for adult 

specimens from the Atlantic and Pacific oceans and the 

Mediterranean Sea were as follows (mm): condylobasal 

length, 472–555 (n 5 66); length of rostrum, 274–343 (n 5 

59); width of rostrum at base, 87–119 (n 5 58); width of 

rostrum at one-half its length, 38–64 (n 5 58); preorbital 

width, 169–196 (n 5 6); postorbital width, 202–239 (n 5 6); 

greatest width of premaxillaries, 75–89 (n 5 10); parietal 

width, 162–184 (n 5 7); length of temporal fossa, 86–119 

(n 5 40); height of temporal fossa, 80–105 (n 5 40); number 

of teeth in each side of upper jaw, 19–26 (n 5 81); and 

number of teeth in each side of lower jaw, 19–28 (n 5 82— 

Miyazaki and Perrin 1994). Cranial measurements obtained 

by 2 of the authors (KLW and JGM) from another 23 adults 

from the Atlantic and the Pacific oceans are almost entirely 

within these ranges except for the following (mm): preorbital 

width, 163–185; postorbital width, 192–215; greatest width 

of premaxillaries, 72–85; and parietal width, 133–160. 

DISTRIBUTION 

Steno bredanensis is found worldwide with reports from 

the Atlantic, Pacific, and Indian oceans, typically in warm 

temperate, subtropical, or tropical waters (Fig. 4). In the 

southwestern Atlantic the most southerly report of the 

species is from Patagonia in Argentina but is based on a 

solitary skull at the Academy of Natural Sciences (Philadelphia, 

Pennsylvania) collected in the late 1800s (catalog 

number ANSP 23360). The species is reported on many 

occasions from Brazil (Pinedo and Castello 1980; Rice 1998; 

Siciliano et al. 2007). In the Caribbean Sea, S. bredanensis is 

reported throughout the Gulf of Mexico and along the 

coastal United States north to Virginia (Miyazaki and Perrin 

1994).


The most northerly records from the northeastern 

Atlantic are from the Scheldt estuary, Belgium, in the North 

Sea (Booij 2004), and from a stranded specimen in Paimpol, 

France (Cuvier 1812; Van Beneden 1889). S. bredanensis also 

is reported from Portugal (Busnel and Dziedzic 1966), Spain 

(Hashmi and Adloff 1991), the Mediterranean Sea (Watkins 

et al. 1987), Macronesia (Bronner et al. 2003; Steiner 1995), 

and the western coast of Africa from Mauritania to Namibia 

(Addink and Smeek 2001; Bronner et al. 2003; Perrin and 

Van Waerebeek 2007; Ross et al. 1985; Steiner 1995). 

A stranding of S. bredanensis is reported off the coast of 

southeastern Africa near the Zambesi River (Best 1971; 

Brownell 1975). However, this may have been a mistake, 

because it appears that this same specimen was later described 

as a junior synonym for Sousa plumbea (Brownell 1975; 

plumbea is currently considered a synonym of Sousa 

chinensis). S. bredanensis is reported from Tanzania and 

Kenya, but not enough detail is provided to evaluate the 

validity of this report (Davies and Vanden Berghe 1994). It is 

also found offshore of Somalia, the Aden District, and the 

Red Sea (Anderson 1891; Ballance et al. 1996; Best 1971; 

Frazier et al. 1987). The species is found in the Arabian Sea 

from the Gulf of Oman throughout the Indian Ocean to the 

Nicobar Islands (Alling 1986; Anderson 1891; Anderson et al. 

1999; R. M. Baldwin et al., in litt.; Ballance et al. 1996, 2001; 

Ballance and Pitman 1998; International Whaling Commission 

1994; Leatherwood and Reeves 1989; Van Waerebeek 

et al. 1999). Strandings are reported from Java, 

Indonesia (Chasen 1940; Kahn 2001), throughout the South 

China Sea (Beasley and Jefferson 1997; Chantrapornsyl 

et al. 1996; Corkeron et al. 2003; Heaney et al. 1998; 

Parsons 1998; Smith et al. 1997; Yang 1976), to the East 

China Sea (Wenji 1980). 

In the western Pacific there are reports from Japan, the 

Northern Mariana Islands, and Kiribati (Hobbs and Jones 

1993; International Whaling Commission 1994; Jefferson 

et al. 2006; Miller 2006; Miyazaki 1980). A specimen also 

was obtained from the Marshall Islands (Smithsonian 

Cetacean Distributional Database, available at http://www. 

cms.int/reports/WAFCET/WAFCET2/WAFCET2_Report. 

htm, accessed 25 May 2011). 

In the central North Pacific, S. bredanensis is commonly 

sighted throughout the Hawaiian Archipelago (Baird et al. 

2008). In the eastern Pacific this species is described as part of 

the fauna in the Bering Sea; however, this report is not 

substantiated by a positively identified specimen (Collins 

et al. 1945). The most northerly reports on the west coast 

of the United States are from Washington and Oregon, but 

these strandings are thought to represent vagrants outside of 

normal species range (Ferrero 1994; Norman et al. 2004). 

S. bredanensis is reported from California (Daugherty and 

Schuyler 1979; Woodhouse 1991), down the coast of Mexico 

(Estrella 1994; Heyning 1986; Perrin and Oliver 1982; 

Smithsonian Cetacean Distributional Database [http://www. 

cms.int/reports/WAFCET/WAFCET2/WAFCET2_Report. 

htm, accessed 25 May 2011]) through Central America (Perrin 

and Kashiwada 1989), from the Pacific coast of Colombia 

(Holt and Jackson 1987; Mora-Pinto et al. 1995; Rodriguez 

1989), and from the Galapagos Islands (Orr 1965). The most 

southerly report along the South American coast is from 

Chile at 24uS (Van Waerebeek and Guerra 1988). S. 

bredanensis is commonly sighted in the eastern tropical Pacific 

but not in high abundance (Au and Perryman 1985; Hewitt 

1985; Wade and Gerrodette 1993; Wahlen et al. 1986). S. 

bredanensis is relatively abundant in the Society Islands, 

French Polynesia, and is occasionally sighted about 1,440 km 

to the north in the Marquesas, French Polynesia (Gannier 

2000, 2002; Gannier and West 2005; Laran and Gannier 

2001). There are also reports from New Zealand waters 

(Baker 1983), the Solomon Islands, Papua New Guinea 

(Brownell et al. 1978), and Australia (Baker 1983; Bannister 

et al. 1996; Ogawa 1938). 

FOSSIL RECORD 

Steno fossils are uncommon but they are reported from 

the lower and middle Pliocene in Europe (Marcuzzi and 

Pilleri 1971). Fossil records for cetacean history extend back 

50 million years and odontocetes most likely originated 

approximately 34–35 million years ago (Fordyce 2002). 

Fossil records for the Delphinidae date back to the late 

Miocene, possibly 11 million years ago, in both Europe and 

North America (Barnes et al. 1985; Fordyce 2002). 

FORM AND FUNCTION 

Form.—The vertebral formula of Steno bredanensis is 7 

C, 13 T, 15–16 L, 30–31 Ca, total 65–67, with atlas and axis 

fused (Miyazaki and Perrin 1994; Tinker 1988). A reduced 

number of vertebrae is reported for 1 adult specimen with a 

vertebral formula of 7 C, 12 T, 15 L, 28 Ca, total 62 (Buchholtz 

and Shur 2004; Buchholtz et al. 2005). There are 21 or 22 

chevron bones, and phalangeal count is I (3), II (8–9), III (6–7), 

IV (3), and V (2—Tinker 1988). We have observed 0–1 floating 

ribs, 6–7 single-headed ribs, and 5–7 double-headed ribs. 

Much of what is known of the anatomy comes from a 

monograph by Neuville (1928). He described the soft 

anatomy of the thyroid, thymus, lymphatic system, lungs, 

stomach, spleen, and pancreas. The liver of S. bredanensis is 

bell shaped with 2 surfaces, a convex facies diaphragmatica 

and a facies visceralis that is divided into 2 lobes (Hojo and 

Mitsuhashi 1975). There is coronal duplication in teeth 

(Neuville 1928). Emmetropia, a state of perfect vision where 

the eye is relaxed while focused on distant objects, and 

movability of the operculum were observed in air for captive 

S. bredanensis (Dral and Duok van Heel 1974). A diagram of 

the tympanic bones is provided in Pilleri et al. (1989). The


nasal sac system is described as similar to that of other 

delphinids (Mead 1975; Purves 1966). 

Masses of selected organs are available from Japanese 

specimens that ranged between 112.4 and 154.7 kg in total 

body weight: brain, 1,170–1,602 g (n 5 14); heart, 605– 

1,080 g (n 5 15); lungs, 1,640–3,934 g (n 5 15); liver, 1,720– 

3,432 g (n 5 15); left kidney, 489–826 g (n 5 14); right 

kidney, 500–830 g (n 5 15); spleen, 34–178 g (n 5 15); 

pancreas, 110–191 g (n 5 14); and intestines, 1,922–3,480 g 

(n 5 15—Miyazaki and Perrin 1994). 

Function.—A physiological study of a captive Steno 

bredanensis compared renal function at 4 and 18 h after a 

meal. At 4 h after a meal, a higher urine flow rate (2.8 ml/ 

min at 4 h compared to 1.3 ml/min at 18 h), glomerular 

filtration rate (169 ml/min at 4 h compared to 131 ml/min at 

18 h), and urea concentration (82.4 ml/min at 4 h compared 

to 53.1 total body weight at 18 h) were observed (Malvin and 

Rayner 1968). Body temperature of an individual in Hawaii 

monitored with telemetry over 29 hours was 36.5–37.6uC 

and mean respiration rate was 2.8 breaths per minute 

(Whittow et al. 1978). 

Hemoglobin of S. bredanensis is categorized as type V, 

having 2 distinct hemoglobin bands (Baluda et al. 1972). 

Blood chemistry and hematology parameters from 17 S. 

bredanensis were compared according to health status (sick 

versus healthy). Sick dolphins have higher levels of aspartate 

aminotransferase, alanine aminotransferase, lactate dehydrogenase, 

bicarbonate, and globulins than healthy ones, 

whereas healthy individuals have higher levels of alkaline 

phosphatase and protein than sick S. bredanensis (Baluda 

et al. 1972). Total white blood cell counts are lower in 

healthy than in sick S. bredanensis, and averaged 7.57 3 10 3 / 

mm 3 in 157 samples collected from 8 healthy individuals 

(C. A. Manire et al., in litt.). A stranded calf in Brazil that 

died had a white blood cell count of 4.4 3 10 3 /mm 3 and low 

values of aspartate aminotransferase, alanine aminotransferase, 

creatinine, and glucose (Bastos et al. 2003). Hematocrit 

from a single individual is reported at 50% (Malvin and 

Rayner 1968) and averaged 47.8% in 170 samples collected 

from 8 healthy individuals (C. A. Manire et al., in litt.). 

Immunoglobulin classes also have been identified from sera 

(Nash and Mach 1971). 

ONTOGENY AND REPRODUCTION 

Length at birth is probably about 100 cm; the largest 

reported fetus measured 93 cm and a stranded calf measured 

106 cm (Bastos et al. 2003; Miyazaki and Perrin 1994; West 

2002). Growth is rapid during the first 5 years of life, with 

individuals reaching at least 200 cm by 5 years of age 

(Miyazaki and Perrin 1994; West 2002). Asymptotic body 

lengths are reached at about 231–258 cm, depending on 

geographic region. In Japan, Steno bredanensis reaches 

asymptotic lengths at 231 cm and in Florida at 239 cm 

(Miyazaki 1980; M. K. Stolen, in litt.). We observed 

asymptotic lengths of 210–231 cm (n 5 62) in a group of 

specimens from the Atlantic and Pacific oceans. In Brazil, 

asymptotic length is greater, estimated at 258 cm (n 5 14— 

Siciliano et al. 2007). 

In Brazil, a large S. bredanensis aged at 6 years old was 

pregnant (Siciliano et al. 2007). Females sampled from the 

Atlantic and Pacific oceans begin a gradual process of sexual 

maturation between 3 and 6 years of age (with age 

determined from undecalcified dentinal layers) and reach 

sexual maturity by 8–9 years (West 2002). Females attain 

sexual maturity at a body length of 212–219 cm and a body 

mass of 101–108 kg (West 2002). Our observations confirm 

data suggesting that females reach sexual maturity at about 

10 years of age and at 210 cm in length in Japan 

(unpublished data cited in Miyazaki and Perrin 1994). 

Males sampled from the Atlantic and Pacific reach 

sexual maturity between 5 and 10 years of age, at a body 

length of 211–216 cm and a body mass of 83–102 kg (West 

2002). A stranded male estimated at 7 years of age was 

considered immature because of small testes mass of 23.3 g 

(Ferrero 1994), and 2 males 14 and 21 years of age 

(determined from both decalcified and undecalcified dentinal 

layers) were sexually mature with testes that contained 

sperm (Miyazaki 1980). Immature males have testes masses 

# 54.3 g, whereas sperm-producing testes of mature males 

have masses of 66–1,500 g (Miyazaki 1980; West 2002). 

Females generally attain physical maturity at a younger 

age and smaller size than males. In females sampled from the 

Atlantic and Pacific, physical maturity, as defined by fusion 

of the vertebral epiphyses, is attained at about 9–12 years of 

age, at a body length of 210–217 cm, and a body mass of 85– 

101 kg (n 5 13—West 2002). Males attain physical maturity 

(vertebral epiphyseal fusion) at a wide range of ages. One 

male was physically mature at only 5 years of age, whereas a 

14-year-old individual was still immature (in a sample of 15 

males—West 2002). A previous report suggested epiphyseal 

fusion in males at 16 years of age (Miyazaki and Perrin 

1994). Males sampled from the Atlantic and Pacific reach 

physical maturity at 227–231 cm and 119–130 kg (West 

2002). No information is available regarding reproductive 

seasonality or the duration of gestation or lactation. 

ECOLOGY 

Steno bredanensis may be solitary but is often found in 

groups of various sizes. The largest report of group size was 

estimated at 160 individuals in the Mediterranean (Watkins 

et al. 1987). Group size averaged 10.8 individuals in French 

Polynesia where solitary sightings and group sizes between 2 

and 35 individuals were observed (Gannier and West 2005). 

Group size averaged 10 individuals from 44 sightings in 

Hawaii with a range of 2–90 dolphins (Baird et al. 2008). In 

the Canary Islands, groups are most commonly composed of


10–20 animals (X¯ 5 16.8), although groups of up to 50 

individuals are sighted (Ritter 2002). 

Large-scale vessel surveys in the eastern tropical Pacific 

estimate the abundance of S. bredanensis at 145,900 

(coefficient of variation [CV] 5 0.32—Wade and Gerrodette 

1993). Vessel surveys conducted in both oceanic waters 

and off the outer continental shelf in the northern Gulf of 

Mexico estimate 2,223 individuals (CV 5 0.41—Fulling et 

al. 2003; Mullin and Fulling 2004; Waring et al. 2005). J. R. 

Mobley (in litt.) conducted aerial surveys within 25 nautical 

miles of the main Hawaiian Islands and estimated a 

population size of 123 individuals (CV 5 0.63). This is an 

underestimate of population size because at least 337 distinct 

individuals were photographically identified over a 6-year 

period throughout the Hawaiian Islands. However, this 

study still suggests a relatively small population size as well 

as site fidelity because of frequent within and between year 

resightings (Baird et al. 2008). The estimated abundance of 

S. bredanensis throughout the entire Exclusive Economic 

Zone of Hawaii, including waters . 25 nautical miles from 

the main Hawaiian Islands, and those in the northwestern 

Hawaiian Islands is 8,709 individuals (CV 5 0.45—Barlow 

2006). It is not known whether the animals occurring around 

Hawaii are part of the same stock as those in the eastern 

tropical Pacific. 

Steno bredanensis is commonly found in shallow 

nearshore, deep offshore, and oceanic waters. There are 

many reports from shallow waters of coastal Brazil and 

Honduras (Carvalho Flores and Ximenez 1997; Kuczaj and 

Yeater 2007; Lodi 1992). In French Polynesia, this species is 

most frequently sighted in 1,000- to 2,000-m depths, 1.8– 

5.5 km from shore (Gannier and West 2005). Similarly in 

Hawaii, sightings are most common in depths . 1,500 m 

and . 3 km from shore (Baird et al. 2008). In the Canary 

Islands, mean sighting depth is slightly . 500 m and mean 

distance is 4.4 km from shore (Ritter 2002). In French 

Polynesia, Hawaii, and the Canary Islands S. bredanensis 

has been identified as relatively abundant. All of these 

locations are characterized by steep volcanic islands that are 

surrounded by deep oceanic waters close to the coast. 

Steno bredanensis is reported to form mixed schools with 

bottlenose dolphins, melon-headed whales (Peponocephala 

electra), false killer whales (Pseudorca crassidens), Fraser’s 

dolphins (Lagenodelphis hosei), humpback whales (Megaptera 

novaeangliae), short-finned pilot whales (Globicephala macrorhynchus), 

spinner dolphins, and Atlantic spotted dolphins 

(Stenella frontalis—Baird et al. 2008; Gannier and West 2005; 

Leatherwood and Reeves 1983; Perrin and Walker 1975; 

Ritter 2002; Scott and Chivers 1990; Watson 1981). 

There are no confirmed reports of predation but it is 

likely, based on anecdotal evidence, that sharks and killer 

whales (Orcinus orca) prey on S. bredanensis. An attack by a 

group of sharks is theorized as the reason why an individual 

beached itself in Texas (Schmidly and Melcher 1974). In 

Hawaii, a captive S. bredanensis left trainers in the midst of 

open ocean experiments when several small sharks, including 

a 4-m shark, were observed in the area (Norris et al. 1965). 

Scarring from bites of the cookie-cutter shark (Isistius 

brasiliensis) is common (Addink and Smeenk 2001; Baker 

1987). 

Diet has been inferred from stomach contents or from 

field observations of suspected foraging. In most cases, 

stomach contents are from stranded individuals and may 

not be representative of the diet of healthy S. bredanensis. 

Stomach contents from strandings in Hawaii consisted 

primarily of nearshore species, including silverside fish 

(Pranesus insularum), saury (Cololabis adocetus), houndfish 

(Tylosurus crocodilus), and an unidentified squid (Shallenberger 

1981). We observed head and tail of a houndfish in 

separate stomach samples from this same stranding. 

Stomach contents from 3 individuals stranded off Oregon 

and Washington included top smelt (Atherinops affinis), jack 

smelt (Atherinops californiensis), and 2 species of squid 

(Ommastrephes bartrami and Onychoteuthis borealijaponia— 

Ferrero 1994). Stomach contents of stranded specimens in 

Florida include the blanket octopus (Tremoctopus violaceus) 

and, in a few cases, a significant amount of algae (Sargassum 

filipendula—Layne 1965). The slender inshore squid (Loligo 

plei) was reported as the primary cephalopod prey from 

Brazilian specimens (dos Santos 2001). A cutlass fish 

(Trichiurus lepturus) and an unidentified bone fish also were 

reported among solid food remains from Brazil (Di 

Beneditto et al. 2001; Ott and Danilewicz 1996). Remains 

include mollusks and fish from the eastern tropical Pacific 

(Perrin and Walker 1975). 

Foraging by S. bredanensis has been observed in the 

field on a number of occasions. In the Azores underwater 

snipefish (Macrorhamphosus scolopax) were fed upon 

simultaneously by both S. bredanensis and Cory’s shearwaters 

(Calonectris diomedea borealis—Steiner 1995). The 

dolphins were observed taking turns feeding from an 

aggregation of the snipefish that formed a ball about 0.5 m 

in diameter. However, no dolphins were observed keeping 

the fish ball together (Steiner 1995). In the southwestern 

Atlantic off the Brazilian coast, in the Abrolhos Bank 

breeding ground, S. bredanensis was observed catching a 

diskfish (Echeneidae—Wedekin et al. 2004). In both Hawaii 

and in the eastern tropical Pacific adult-sized mahi-mahi 

(Coryphaena hippurus) were reported in the mouth of 

S. bredanensis (Brower 1979; Pitman and Stinchcomb 

2002). We observed mahi-mahi among the stomach contents 

of a specimen incidentally caught in the eastern tropical 

Pacific. 

Helminths described in S. bredanensis include cestodes, 

nematodes, trematodes, and acanthocephalans. Cestodes 

include Tetrabothrius forsteri, Strobilocephalus triangularis, 

and Trigonocotyle prudhoei (Arvy 1982; Delyamure 1955; 

Forrester and Robertson 1975; Gibson and Harris 1979;


Layne 1965). We observed the cestodes Phyllobothrium and 

Monorhygma in stranded specimens from both Hawaii 

and Virginia. Trematodes found in S. bredanensis include 

Campula palliata, Pholeter gastrophilus, Synthesium tursionis, 

and Braunina cordiformis (Forrester and Robertson 

1975; Ott and Danilewicz 1996), and we observed trematodes 

of the genus Nasitrema. Nematodes reported from S. 

bredanensis include Anisakis and Crassicauda (Forrester and 

Robertson 1975). The only acanthocephalan reported in S. 

bredanensis is Bolbosoma capitatum (Gibson and Harris 

1979). An isopod ectoparasite of a fish, Nerocila, was found 

in the stomach of a stranded S. bredanensis in southern 

Brazil (Ott and Danilewicz 1996). 

Solitary and mass strandings have occurred in most 

regions of the world where this species is found. In a mass 

stranding in Cap Vert, Senegal, 25 of 28 animals were males, 

indicating an extremely unbalanced sex ratio (Cadenat 

1949). We examined specimens at the National Museum of 

Natural History, Washington, D.C., from a mass stranding 

in 1976 of 17 individuals from Hawaii and 16 from Virginia 

Beach, Virginia, that had beached later that same year. In 

2002, 14 animals were found dead at Wreck Island, Virginia 

(West 2002). Historical and recent mass strandings have 

been reported for Florida, including 16 near Rock Island in 

1961 (Layne 1965). Recent strandings have occurred along 

both the Gulf of Mexico and the Atlantic coasts, as well as in 

the Florida Keys (C. A. Manire, in litt.; H. L. Rhinehart 

et al., in litt.). Chronic progressive viral arteritis is a 

suspected cause behind some of the Florida strandings 

(H. L. Rhinehart et al., in litt.). 

HUSBANDRY 

Steno bredanensis has been held in captive facilities 

around the world, including in the United States, Hong 

Kong, Japan, Europe, and French Polynesia (Collet 1984; 

Miyazaki 1980; Parsons 1998; Reeves and Leatherwood 

1984; West 2002). In the 1950s, S. bredanensis was captured 

in the Mediterranean Sea and near the Madeira Archipelago, 

Portugal, for a physiological laboratory (Collet 1984). 

In the 1960s and 1970s in Japan, individuals of this species 

were captured and held in a captive facility for at least 

4 months (Miyazaki 1980; Nishiwaki 1967). As of 1981, 23 

individuals had been collected near Hawaii and held at Sea 

Life Park (Waimanalo, Hawaii). Five of these either escaped 

or were released. One of the individuals had initially 

stranded and then survived in captivity for at least 6 years 

(Reeves and Leatherwood 1984; Tomich 1986). In 1971, a 

viable hybrid of a bottlenose dolphin and S. bredanensis was 

born at Sea Life Park, Hawaii (Dohl et al. 1974). At least 5 

S. bredanensis were held in captivity on the island of Moorea 

in French Polynesia for up to 6 years during the 1990s. One 

was a stranded calf that lived for at least 3 years in captivity 

(West 2002). S. bredanensis also has been rehabilitated in 

Florida, where 14 individuals were treated following several 

separate stranding events (C. A. Manire et al., in litt.). A 

stranded specimen in China was cared for at Ocean Park 

Hong Kong (R. E. Kinoshita; in litt.). 

BEHAVIOR 

Steno bredanensis commonly forms subgroups that may 

be a part of a larger group (Kuczaj and Yeater 2007; Ritter 

2002). These tight but dynamic subgroups are often composed 

of 2–8 individuals involved in synchronous behavior. 

The social organization may be of fission–fusion type 

(Kuczaj and Yeater 2007; Ritter 2002). Association patterns 

have demonstrated strong social bonds, especially between 

mothers and calves or juveniles (Kuczaj and Yeater 2007). 

Skimming along at the surface is a common behavior 

(Jefferson 2002; Jefferson and Leatherwood 1993; Leatherwood 

and Reeves 1983). Surface swimming has been 

categorized in Honduras and the Canary Islands as traveling 

(21–27%), resting (4–10%), or milling (9–24%—Kuczaj and 

Yeater 2007; Ritter 2002). Play with objects (7%) and sexual 

behavior (4%) were less commonly observed (Kuczaj and 

Yeater 2007). In other locations, S. bredanensis is known for 

a propensity to associate or play with marine life or flotsam 

such as logs, plywood, and plastic bags (Brower 1979; 

Jefferson 2002; Leatherwood et al. 1982; Pitman and 

Stinchcomb 2002; Ritter 2002; Steiner 1995). Reports of 

cooperative feeding among group members also are common; 

they have been observed engaging in cooperative fishherding 

techniques (Addink and Smeenk 2001) and appeared 

to take turns approaching a small aggregation of 

snipefish (Steiner 1995). Group feeding has been frequently 

observed in the Canary Islands and Honduras (Kuczaj and 

Yeater 2007; Ritter 2002). 

Captive individuals of S. bredanensis have earned a 

reputation as being highly trainable, creative, sociable, and 

easy for humans to work with (Pryor et al. 1969; Tomich 

1969). Many of the behaviors elicited by individuals at Sea 

Life Park (Waimanalo, Hawaii) were new and unreinforced 

behaviors that led to complex training sessions (Pryor et al. 

1969). 

Diving capabilities may be more extreme than observed. 

S. bredanensis is reported to dive as deep as 70 m, where 

a few individuals rubbed against a hydrophone at this 

depth (Watkins et al. 1987). The deepest dive of a trained 

S. bredanensis in open water is 30 m. However, this training 

session ended because the animal became agitated due to the 

presence of sharks (Norris et al. 1965). This specific 

individual was able to dive frequently, because it made 51 

dives in , 2 h during open-water experiments (Norris et al. 

1965). S. bredanensis may remain submerged for up to 15 min 

(Jefferson 2002; Miyazaki and Perrin 1994). While bowriding 

(riding the waves produced off the bow or front of a 

moving vessel) S. bredanensis has attained swim speeds of up


to 16 km/h (Watkins et al. 1987). Although described as only 

an occasional bow-rider (Leatherwood and Reeves 1983; 

Leatherwood et al. 1982), there are reports of bow-riding 

from the Mediterranean Sea, the Azores, the Canary Islands, 

French Polynesia, Hawaii, and Honduras (Baird et al. 2008; 

Kuczaj and Yeater 2007; Ritter 2002; Steiner 1995; Watkins 

et al. 1987; West 2002). 

Some observations of mothers, calves, or mother–calf 

pairs of S. bredanensis have been noted from various regions 

of the world. Twenty-nine percent of the groups sighted in 

French Polynesia and 16% of those sighted in the Canary 

Islands include at least 1 calf (Gannier and West 2005; Ritter 

2002). Echelon swimming occurs, where calves swim very close 

to the dorsal fin of the mother and gain a hydrodynamic boost 

(Addink and Smeenk 2001). Off the coast of Mauritania, in 

North Africa, a mother provided her calf with assistance 

during feeding by encouraging the calf to pick up a fish it had 

dropped (Addink and Smeenk 2001). A stranded calf of S. 

bredanensis in Moorea, French Polynesia, induced lactation in 

an adult bottlenose dolphin held in the same lagoon (Gaspar 

et al. 2000). Epimeletic behavior (care-giving behavior by a 

healthy individual to a compromised or dead individual) has 

been described in Brazil and the Canary Islands (Lodi 1992; 

Ritter 2007). In the Canary Islands, a large adult female, which 

was presumed to be the mother, supported a dead calf at the 

surface. This mother and dead calf were resighted over several 

days, and 2 other adults also were observed providing support 

to the body of the calf (Ritter 2007). 

At sea S. bredanensis produces extremely brief echolocation 

clicks in bursts of only 0.1–0.2 s in duration (Watkins et al. 

1987). Echolocation clicks have both low- and high-frequency 

components ranging between 2.7 kHz and 256 kHz. The 

maximum peak frequency of echolocation clicks is approximately 

25 kHz (Norris 1969). This value is low compared to 

maximum peak frequencies that have been described for other 

species of delphinids such as the false killer whale (40 kHz), the 

Risso’s dolphin (Grampus griseus—50 kHz), the spinner 

dolphin (60 kHz), and the bottlenose dolphin (110–130 kHz— 

Au 2000; Ketten 1984; Madsen et al. 2004). Whistles have been 

recorded with sound frequencies of 3–12 kHz where sequences 

are approximately 0.5 s in duration (W. E. Evans, in litt.; 

Watkins et al. 1987). S. bredanensis has emitted stereotyped 

calls during stressful situations (W. E. Evans, in litt.). Based on 

recordings from 6 sightings, other individuals of S. bredanensis 

remain silent while a member of the group echolocates, 

suggestive of eavesdropping (Gotz et al. 2005). 

GENETICS 

Both mitochondrial and nuclear variation indicate that 

Steno bredanensis is a highly distinctive species. Studies using 

mitochondrial cytochrome-b sequences indicate that S. 

bredanensis differs considerably from other closely related 

species, a finding consistent with placement of this species in 

a separate genus (Agnarsson and May-Collado 2008; LeDuc 

et al. 1999; May-Collado and Agnarsson 2006). Steno and 

Sotalia are sister taxa, according to Bayesian phylogenetic, 

maximum-parsimony, maximum-likelihood, and neighborjoining 

analyses of cytochrome-b sequence variation within 

family Delphinidae, and they are currently retained in the 

subfamily Steninae (Agnarsson and May-Collado 2008; 

LeDuc et al. 1999; May-Collado and Agnarsson 2006). 

However, recent relationships described from a combination 

of mitochondrial and nuclear DNA phylogenies suggest that 

Steno and Sotalia are not sister taxa and that Steninae may 

be an artificial grouping. Instead, the combination of 

mitochondrial and nuclear DNA indicate that Steno should 

be allied with the subfamily Globicephalinae or retained as 

the sole member of Steninae, and that further study is 

needed to resolve this issue (Caballero et al. 2008). 

Isoenzyme analysis of protein variation indicates a low 

proportion of polymorphic loci (5.3%) from an analysis of 

19 loci from 12 enzymes in a population of S. bredanensis in 

waters off Taiji, Japan (Shimura and Numachi 1987). 

Average heterozygosity of this same population was 0.007 

6 0.024 SE, which was the lowest of 12 species of odontocetes 

examined (Shimura and Numachi 1987). Within family Delphinidae, 

average coefficient of genetic distance for isoenzyme 

analyses between S. bredanensis and 7 other delphinids is 0.379, 

compared to the distance of 1.004 between S. bredanensis and 

an average of 4 species of phocoenids (Shimura and Numachi 

1987). Differentiation among microsatellite results indicates 

distinct populations of S. bredanensis at 2 different island 

groups in French Polynesia, which supports site fidelity in this 

region (Oremus 2008). 

CONSERVATION 

Population size and status of Steno bredanensis is poorly 

known from most regions of the world. The Convention on 

International Trade in Endangered Species of Wild Fauna and 

Flora (2011) lists S. bredanensis in Appendix II, as a species 

that is not necessarily now threatened with extinction but 

that may become so unless trade is closely controlled. The 

International Union for Conservation of Nature and Natural 

Resources (2011) considers S. bredanensis a species of ‘‘Least 

Concern.’’ Despite this, a number of potential threats have 

been identified, including fisheries that target S. bredanensis, 

the occurrence of this dolphin as incidental catch, and other 

types of fishery interactions. Additionally, S. bredanensis has 

been live-captured in small numbers for public display, and it 

may be vulnerable to the negative effects of contaminant 

burdens and oceanic debris (Kucklick et al. 2002; Leatherwood 

and Reeves 1989; Miyazaki 1983; Nishiwaki 1967; Oliveira de 

Meirelles and Duarte do Rego Barros 2007; Schlais 1984). 

Fisheries that target S. bredanensis directly are reported 

from Japan, the Solomon Islands, Papua New Guinea, Sri 

Lanka, the Caribbean, and West Africa. In Japan, schools of


S. bredanensis in nearshore waters have been targeted by drive 

fisheries (those that drive the dolphins into shore and kill 

them—Nishiwaki 1967). S. bredanensis is occasionally present 

in the fish market in Shiogama, Japan (Whitehead et al. 2000). 

A group of 23 animals was taken in an Okinawan drive fishery 

in 1976, but none were taken between the years of 1977 and 

1981 (Miyazaki 1983; Nishiwaki and Uchida 1977). In the 

calendar year 1985, Japanese drive fisheries reported a take 

of 60 S. bredanensis throughout Japan (Anonymous 1987). 

Although drives of dolphins are reported as rare events, when 

they do occur, tens of animals may be killed in a single drive 

(Whitehead et al. 2000). Similarly, native fishermen in the 

Solomon Islands and Papua New Guinea target entire schools 

of S. bredanensis in drive fisheries (Takekawa 1996; Young 

and Iudicello 2007). Various dolphin species, including S. 

bredanensis, are caught in Sri Lanka and sold as either 

dolphin or ‘‘dugong’’ meat in local fish markets (Leatherwood 

and Reeves 1989). At least 7 S. bredanensis were 

confirmed at the main fish market in Trincomalee, Sri Lanka, 

between 1983 and 1986 (Leatherwood and Reeves 1989). On 

the Caribbean island of St. Vincent, dolphin fishermen pursue 

all odontocete species encountered, including S. bredanensis, 

but the total take is unknown (Caldwell and Caldwell 1975; 

Van Bree 1975). A small fishery that targets S. bredanensis 

was reported from the western coast of Africa (Mitchell 1975). 

Local fishermen from the Atlantic island of St. Helena have 

harpooned S. bredanensis, but this activity is rare, because the 

animals are known to be difficult to subdue (Perrin 1985). 

Steno bredanensis has been indirectly taken as incidental 

catch from several locations around the world. These include 

coastal and offshore Japan (drift-net fishery), Sri Lanka 

(gill-net fishery), the eastern tropical Pacific (purse-seine 

fishery), Brazil (gill-net fishery), and the Mediterranean Sea 

(gill-net fishery—Alling 1986; Anonymous 1987; Hobbs and 

Jones 1993; International Whaling Commission 1994). The 

annual number of S. bredanensis incidentally caught from 

any of these locations is generally very low, with the highest 

report being 18 animals caught in the high seas of Japan in 

1990 (Hobbs and Jones 1993; International Whaling 

Commission 1994). Despite only 6 animals being reported 

from Brazilian gill-net fisheries in 1990, the magnitude of 

bycatch for S. bredanensis is considered higher than for other 

small cetaceans in Brazil (International Whaling Commission 

1994; Monteiro-Neto et al. 2000; Siciliano 1994). 

Steno bredanensis has been captured and subsequently 

held captive in facilities around the world. However, there 

are no reports of recent collections of this species for 

captivity, and it is unknown if past collections had a 

significant impact on local populations. 

Bait stealing by S. bredanensis has been reported as a 

problem for fisheries in Hawaii and may also occur in other 

regions of the world (Schlais 1984). Fishermen have reported 

losing 20–50% of bait they set, and this species is blamed for 

all offshore losses in Kona, Hawaii (B. A. Kuljis, in litt.). 

There are anecdotal reports of illegal activities (poisoning 

and shooting) by local fishermen to mitigate their losses. 

This may have an effect on population numbers of S. 

bredanensis in Hawaiian waters (Schlais 1984). 

Chemical contamination of the oceans is a widespread 

problem and may be particularly threatening to high-level 

consumers such as dolphins. Low concentrations of dichlorodiphenyltrichloroethane 

(DDT), dieldrin and trans-nonachlor, 

and polychlorinated biphenyl levels up to 39 ppm were detected 

in the blubber, brain, or muscle tissues in 6 of 7 S. bredanensis 

that mass-stranded in Hawaii (O’Shea et al. 1980). In the Gulf of 

Mexico, levels of persistent organic pollutants in S. bredanensis 

were similar to or lower than those reported from other 

odontocetes (Kucklick et al. 2002; Struntz et al. 2004). However, 

S. bredanensis hashigherpercentageratiosof4,49-DDT to 4,49- 

DDT + 4,49-dichlorodiphenyldichloroethylene (4,49-DDE) 

when compared to those measured in bottlenose dolphins 

(Kucklick et al. 2002; Struntz et al. 2004). High concentrations 

of contaminant persistent organic pollutants found in immature 

dolphins suggest an off-loading of the contaminant from 

mothers to calves (Kucklick et al. 2002). 

Oceanic debris represents an additional threat to S. 

bredanensis. An emaciated individual that was live-stranded 

in Brazil had ingested 2 plastic bags (Oliveira de Meirelles 

and Duarte do Rego Barros 2007). Plastic bags also have 

been found in the stomachs of stranded individuals from 

Hawaii and Virginia (Walker and Coe 1990). 

ACKNOWLEDGMENTS 

We are grateful to C. Potter and A. Heilman for 

assisting with skull measurements. We would like to 

acknowledge S. Courbis and A. Brittain for assistance with 

the genetics section and A. Pacini for assistance with the 

echolocation section. We also express thanks to M. 

Kuprijanova and R. Brownell. 

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Associate editor of this account was BARBARA 

MEREDITH J. HAMILTON. 

BLAKE. Editor was

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