Geographic Range
Puffinus auricularis
has a disjunct range, with two subspecies occurring in different parts of the Pacific
Ocean and associated breeding islands. Some authorities recognize these two subspecies
as separate species based on their range differences and some morphological and ecological
differences.
Townsend's shearwaters (
Puffinus auricularis townsendi
) are found on breeding colonies in the Revillagigedo Islands, off the west coast
of Mexico. They were previously known to breed on 3 islands: Clarion, San Benedicto,
and Socorro, but are now only found breeding on Socorro Island. When not at breeding
colonies they are found over oceanic waters over the continental shelf off the western
coast of Mexico and Central America.
Newell's shearwaters (
Puffinus auricularis newelli
) are found in breeding colonies on the Hawaiian Islands. They once bred on islands
throughout the Hawaiian archipelago, but are now found on Moloka'i, Kaua'i, and Hawai'i
and possibly O'ahu, Maui, Lana'i, and Lehua Islet. When not at breeding colonies they
are found over deep, tropical oceanic waters south and west of the Hawaiian Islands.
- Biogeographic Regions
- oceanic islands
- pacific ocean
Habitat
Townsend's and Newell's shearwaters are pelagic birds that breed at higher elevations
on oceanic islands. These shearwaters cannot take flight from the ground so must take
advantage of either elevation to take flight or must climb a tree or rock outcrop
to gain sufficient height to take flight. The two subspecies of
Puffinus auricularis
differ in their pelagic habitats, but both avoid nearshore waters generally, except
when they form large evening rafts before returning to breeding colonies.
Townsend's shearwaters
breed in rocky burrows in shrubby or grassy areas, typically at elevations above
500 m on Socorro Island. They bred at elevations above 244 m in Clarion Island and
above 161 m on San Benedicto Island. These islands are arid and the vegetation tends
to be shrubby. Townsend's shearwaters are found over cold, continental slope waters
(depths of 200 to 2000 m) when not at breeding colonies.
Newell's shearwaters
also breed in burrows, typically in areas where the roots of
Metrosideros polymorpha
trees form the burrows and there is a covering of uluhe fern (
Dicranopteris linearis
), although burrows in tussock-grass habitats also occur. They breed at elevations
from 160 to 1200 meters, although breeding elevation may be influenced by the presence
of wedge-tailed shearwaters (
Puffinus pacificus
), which are found at lower altitudes. Currently, most breeding burrows occur on steep
slopes where non-native terrestrial predators are less likely to be found. Breeding
islands in Hawaii are marked by their extreme mesic vegetation, with some of the highest
annual rainfall amounts worldwide. Newell's shearwaters are found over deep (depths
over 2000 m), tropical and subtropical oceanic waters when not at breeding colonies.
- Habitat Regions
- tropical
- terrestrial
- saltwater or marine
- Terrestrial Biomes
- savanna or grassland
- rainforest
- scrub forest
- Aquatic Biomes
- pelagic
Physical Description
Townsend's and Newell's shearwaters are medium-sized
shearwaters
, with body lengths of about 33 cm. They have black or sooty brown plumage on their
dorsal surfaces and white on ventral surfaces. There is a sharp line from the bill
along the cheek to right behind the head that demarcates dorsal and ventral surfaces.
They have a white side patch that extends on to their rump dorsally on each side.
Ventrally, the wings are marked by a sharp black line on the leading and trailing
edges, with the feathers being white otherwise. The bill is dark gray and sharply
hooked at the tip and the legs are pale pink with a black stripe that runs along the
back edge of the leg and a black line along the outer surface of the toes. The eyes
are brown. The undertail coverts are white at the base and black at their tips in
Newell's shearwaters. Townsend's shearwaters have completely dark undertail coverts.
The two subspecies can be distinguished primarily by their largely non-overlapping
distribution, but also differ in tail length. Townsend's shearwaters have shorter
tails, from 71 to 83 mm (76.3 mm average), Newell's shearwaters have longer tails,
from 77 to 89 mm (83.9 mm average). These shearwaters have relatively high wing-loading,
with short and broad wings typical of diving shearwaters and consistent with their
flight style and non-migratory behavior.
Hatchlings are covered with grey down, with bare lores, foreheads, and chins. The
bill is grey, the eyes are black, and the legs are pink on the inner surface and grey
on the outer surface. The timing of molts is not known, although it seems to occur
once yearly in adults.
There is no reported sexual dimorphism in size or color. Townsend's shearwaters are
from 315 to 358 grams. Newell's shearwaters are from 342.5 to 425 grams, although
these numbers are based on relatively few individuals. Wing lengths are from 220 to
238 mm in Townsend's shearwaters and 223 to 249 mm in Newell's shearwaters.
Townsend's and Newell's shearwaters can be confused with other species of shearwaters.
They are larger than Audubon's shearwaters (
Puffinus lhermineri
) and little shearwaters (
Puffinus assimilis
). They are similar in size to Manx (
Puffinus puffinus
), black-vented (
Puffinus opisthomelas
), and wedge-tailed shearwaters (
Puffinus pacificus
). They are smaller than pink-footed shearwaters (
Puffinus creatopus
).
- Other Physical Features
- endothermic
- homoiothermic
- bilateral symmetry
- Sexual Dimorphism
- sexes alike
Reproduction
Townsend's and Newell's shearwater form pairs in the year before their first breeding
attempt, which is estimated to occur at approximately 6 years old. The sex ratio is
1:1 in these shearwaters and they seem to beed monogamously for life.
- Mating System
- monogamous
Newell's shearwaters return to breeding colonies in April, when mated pairs scout
for nest burrows. Nest burrows are about 87.7 cm long and are re-used yearly by shearwaters.
After nesting sites are established, males and females leave the breeding colony to
forage and replenish nutrient reserves for egg-laying and incubation. Males and females
return to the breeding colony and begin egg-laying and incubation in early June. Egg-laying
is synchronized in colonies and incubation lasts 53-54 days. Hatchlings fledge after
81 to 94 days (average 92.4). Hatchlings weight about 364.6 g just before fledging.
The timing of breeding and egg-laying in Townsend's shearwaters varies slightly from
Newell's shearwaters, but has not been studied as well. Estimates suggest that egg-laying
occurs in January on Socorro Island. The timing of egg-laying in these shearwaters
may vary across breeding colonies as well. Historical data also suggest that breeding
may not be as synchronous as in Newell's shearwaters. Townsend's shearwaters fledge
in less time than do Newell's shearwaters, although those measurements are not reported
in the literature.
Townsend's and Newell's shearwaters begin to form pair bonds at 4 to 5 years old,
presumably the age of sexual maturity.
- Key Reproductive Features
- iteroparous
- seasonal breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- oviparous
Male and female parents care for their young in nest burrows. Both develop incubation
patches shortly after egg-laying. Observations suggest that mates share incubation
responsibilities equally. Males incubate first, when females forage after egg-laying
to replenish their reserves. Time spent incubating is not known and there is a possibility
that eggs are neglected sometimes as parents undertake long commutes to foraging areas.
Young are born in an altricial state and are unable to thermoregulate. Hatchlings
are tended for 5 to 10 days after hatching and are fed at varying intervals. Estimates
based on foraging commute distances and fledging period length suggest that Townsend's
shearwater hatchlings are fed once every 1.2 to 1.3 days and Newell's shearwaters
are less often (fledging periods are 23% longer). As soon as the young fledge, they
become independent.
- Parental Investment
- altricial
-
pre-fertilization
- provisioning
-
protecting
- female
-
pre-hatching/birth
-
provisioning
- female
-
protecting
- male
- female
-
provisioning
-
pre-weaning/fledging
-
provisioning
- male
- female
-
protecting
- male
- female
-
provisioning
Lifespan/Longevity
Longevity of Townsend's and Newell's shearwaters is not reported. Estimates suggest
that annual adult survivorship is 90%.
Behavior
Townsend's and Newell's shearwaters are unusual among other night-active seabirds,
which are generally crepuscular in habits (
dark-rumped petrels
, for example). Townsend's and Newell's shearwaters arrive and depart from breeding
colonies in full dark, arriving well after sunset and departing in the early pre-dawn
hours. Newell's shearwaters were measured flying at from 8 to 800 meters above the
ground. Moon phase doesn't seem to impact nocturnal activity, although the presence
of man-made lighting seems to attract shearwaters on Hawaii, resulting in mortality.
These shearwaters do not seem to be migratory, occurring year round in their pelagic
foraging areas. However, concentrations of adult, breeding birds occurs within several
hundred kilometers of breeding colonies during breeding season in Newell's shearwaters.
At other times of the year these shearwaters are not found near the Hawai'ian islands.
These shearwaters are colonial breeders.
Townsend's and Newell's shearwaters do not walk well because of the placement of their
legs on their bodies. They climb well and must gain altitude before being able to
take flight. Their flight is fast and low to the water. They beat their wings rapidly
and are not generally seen soaring. They have high wing loading because of their short,
rounded wings, presumably influenced by selection towards smaller wings for underwater
swimming. Shearwaters rest often on the water because their flight is relatively energy
expensive. Average flight speed has been estimated at 12.3 m/s into a headwind of
17.6 m/s. Their high wing loading means that landing in breeding colonies often involves
crashing into vegetation and the ground.
Home Range
Townsend's shearwaters were recorded ranging up to 650 km from colonies. Newell's
shearwaters were recorded ranging up to 1300 km from colonies. Nests are spaced 3
to 10 meters apart in Townsend's shearwater colonies and 20 to 50 meters apart in
Newell's shearwater colonies.
Communication and Perception
Townsend's and Newell's shearwater use vocalizations to communicate among themselves.
Adults use an "aerial" call in flight or on the ground near their nesting burrows.
It is characterized as a series of 2 to 6 gruff, braying calls. The native Hawai'ian
name for Newell's shearwaters is "a'o" - which mimics the aerial call. Hatchlings
make a squeakier version of the aerial call, their voices deepen as they age. When
mated pairs are at a burrow together, they duet with aerial calls. Vocalizations in
colonies are generally at night, with birds vocalizing when they arrive and depart.
- Other Communication Modes
- duets
Food Habits
Townsend's and Newell's shearwaters use a pursuit-plunge foraging style. They fly
above the water's surface and dive in when they see prey. They continue swimming with
partly folded wings underwater and can swallow prey while swimming. They are estimated
to reach depths up to 10 meters using this strategy. Prey are not well reported in
the literature and only squid were recorded as part of regurgitates at nests of Newell's
shearwaters. These shearwaters probably eat primarily fish and squid.
Newell's shearwaters are documented foraging at sea along ocean fronts in the Equatorial
Counter Current. Like, Townsend's shearwaters, they concentrate foraging effort on
highly productive areas, which often involves a high investment in travel to foraging
areas, and rest often. Newell's shearwaters are known as "tuna birds," birds that
form multi-species flocks and forage above tuna schools. They seem to rely on the
presence of schools of predatory fish, especially
tunas
, to drive suitable fish prey to near the ocean's surface.
Townsend's shearwaters seem to forage along upwelling fronts of the Costa Rica Current
over the continental shelf along Mexico's west coast. Shearwaters travel far to forage,
but focus their foraging on highly productive areas and rest frequently. Townsend's
shearwaters forage in areas where prey occurs closer to the surface and are not known
to typically associate with schools of tuna. However, they do forage near groups of
foraging
porpoises
, which may also drive prey to nearer the surface.
- Primary Diet
- carnivore
- Animal Foods
- fish
- mollusks
Predation
There are no reports of native predators of Townsend's and Newell's shearwaters. Rarely
a shearwater diving above a school of foraging tuna may be taken by a tuna and their
nocturnal arrival and departure from breeding colonies is thought to be an adaptation
to avoid avian predators. Volcanic eruptions and hurricanes have been cited as the
largest source of non-human influenced mortality. With the invasion of their breeding
islands by non-native mammalian predators, including rats, pigs, mongooses, barn owls,
and domestic cats, mortality rates of adults, eggs, and hatchlings has soared, leading
to severe population declines and the loss of entire breeding populations.
Ecosystem Roles
Townsend's and Newell's shearwaters are rare and are unlikely to have significant
ecosystem impacts, although they are predators of small fish and squid. Ectoparasites
recorded include biting lice (
Mallophaga
) and nest mites (
Ornithodosis capensis
).
Plasmodium
has been found in the blood of Newell's shearwaters, but none have shown signs of
avian malaria. This is true of wedge-tailed shearwaters (
Puffinus pacificus
) in Hawai'i as well, suggesting that these birds are not prone to avian malaria.
These shearwaters may forage in association with tunas (
Thunnus
) and porpoises (
Phocoenidae
) to take advantage of small fish scared to the ocean's surface. Jaegers (
Stercorarius
species) and South Polar skuas (
Catharacta maccormicki
) steal fish from Townsend's and Newell's shearwaters.
- tunas ( Thunnus species)
- porpoises ( Phocoenidae )
- biting lice ( Mallophaga )
- nest mites ( Ornithodosis capensis )
- jaegers ( Stercorarius species)
- South Polar skuas ( Catharacta maccormicki )
Economic Importance for Humans: Positive
Townsend's and Newell's shearwaters are important members of pelagic ecosystems, although
their current numbers suggest they have minimal impact in pelagic food webs. When
they were abundant on the Hawai'ian Islands, native Hawai'ians collected shearwaters
for food.
- Positive Impacts
- food
Economic Importance for Humans: Negative
There are no known adverse effects of Townsend's or Newell's shearwaters on humans. They are found in remote areas and have few direct interactions with humans.
Conservation Status
Puffinus auricularis
is critically endangered as both
P. a. auricularis
and
P. a. newelli
. Newell's shearwaters (
P. a. newelli
) are threatened under the U.S. Endangered Species Act.
Townsend's shearwaters have been extirpated from two breeding islands, Clarión and
San Benedicto islands. Townsend's shearwater populations on Clarión Island were ravaged
by introduced
sheep
,
pig
, and
rabbit
populations, resulting in total extirpation by 1988. Populations on San Benedicto
Island were destroyed by a volcanic eruption in 1952 and have not been recorded breeding
on the island since then, although small numbers are reported on the island. The population
is now limited to a single, small breeding area on Socorro Island where the vegetation
has been seriously degraded by sheep (
Ovis aries
) grazing. Feral cats (
Felis catus
) are also found on this island and are reducing their numbers. Up to 92% of cat scats
on Socorro Island above 500 m contain shearwater remains and it was estimated that
cats take about 350 females yearly. Estimates of the population size on this island
were around 1100 in 2004. These estimates suggest demographic instability and extinction
within 100 years. Sea censuses suggest larger population sizes, as many as 46,000
(but with a range estimate of 18,000 to 89,000, and representing approximately 10,600
breeding birds). The eradication of introduced mammals on Socorro Island is vital
to protecting the population. The Mexican navy has greatly reduced sheep populations,
but rabbit eradication programs have not been successful. Pigs and sheep were eradicated
from Clarión Island in 2002 and taped playback of Townsend's shearwater calls are
being used to attract potential breeding birds.
Newell's shearwaters were thought to be extinct in Hawaii by 1908, but a population
was rediscovered off Kaua'i in 1947. Populations on Hawaii have declined dramatically,
by over 60% in just a 3 year period between 1999 and 2001 and about 72% from 1993
to 2001. Introduced mongooses (
Herpestes javanicus
), rats (
Rattus
), cats (
Felis catus
), and barn owls (
Tyto alba
) all prey on these shearwaters in Hawai'i. Introduced pigs (
Sus scrofa
), goats (
Capra hircus
), and strawberry guava (
Psidium cattleianum
) are resulting in habitat degradation affecting nesting colonies. Newell's shearwaters
are attracted to man-made lights in Hawai'i and this attraction has resulted in high
mortality, especially of fledglings. Shielding light sources so that they don't shine
upwards decreases their attraction to shearwaters. These shearwaters also suffer mortality
as a result of collisions with man-made structures, especially power lines. Hundreds
of shearwaters die from collisions with powerlines each year, the majority are fledglings
and most collisions seem to occur because power lines are located in river valleys
used for traveling from colonies to the ocean or because fledglings are attracted
towards the land by man-made lights.
Sea censuses indicate population sizes of approximately 84,000 (but with an estimate
range of 57,000 to 115,000, and representing approximately 18-19,000 breeding birds).
Vocalization play back and burrow construction are being used to attract birds to
breeding colony sites at Kilauea Point National Wildlife Refuge, where there are few
obstacles and little artificial lighting to interfere with the movement of shearwaters.
All non-essential lights are turned off or shielded in Kauai'i from mid-September
to mid-December, when juveniles are leaving breeding colonies.
Breeding bird populations on San Benedicto Island, previously an important breeding
colony site for Townsend's shearwaters, have changed substantially in recent decades.
Possible changes in local marine ecosystems have resulted in populations of central
Pacific birds starting to breed on this island, including Laysan albatrosses (
Phoebastria immutabilis
), black-footed albatrosses (
Phoebastra nigripes
), red-tailed tropicbirds (
Phaethon rubricauda
), and brown boobies (
Sula leucogaster
).
Other Comments
Townsend's (
P. a. auricularis
) and Newell's shearwaters (
P. a. newelli
) are currently considered subspecies of
P. auricularis
by the American Ornithological Union and the Howard and Moore Checklist of the Birds
of the World (2003). However, Brooke (2004) recognized them as separate species and
various conservation organizations have adopted that recommendation. They have non-overlapping
distributions, vary somewhat ecologically, and are morphologically distinguishable.
Care was taken to distinguish between information on
P. a. auricularis
and
P. a. newelli
in this account to recognize these differences.
Shearwaters were considered an omen of death by native Hawai'ian islanders because
of their long absence from terrestrial breeding colonies each year.
Additional Links
Contributors
Tanya Dewey (author), Animal Diversity Web.
- oceanic islands
-
islands that are not part of continental shelf areas, they are not, and have never been, connected to a continental land mass, most typically these are volcanic islands.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Pacific Ocean
-
body of water between the southern ocean (above 60 degrees south latitude), Australia, Asia, and the western hemisphere. This is the world's largest ocean, covering about 28% of the world's surface.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- tropical
-
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
- terrestrial
-
Living on the ground.
- saltwater or marine
-
mainly lives in oceans, seas, or other bodies of salt water.
- tropical savanna and grassland
-
A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.
- savanna
-
A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.
- temperate grassland
-
A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.
- rainforest
-
rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.
- scrub forest
-
scrub forests develop in areas that experience dry seasons.
- pelagic
-
An aquatic biome consisting of the open ocean, far from land, does not include sea bottom (benthic zone).
- endothermic
-
animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.
- bilateral symmetry
-
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
- monogamous
-
Having one mate at a time.
- iteroparous
-
offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).
- seasonal breeding
-
breeding is confined to a particular season
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- oviparous
-
reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.
- altricial
-
young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.
- diurnal
-
- active during the day, 2. lasting for one day.
- crepuscular
-
active at dawn and dusk
- motile
-
having the capacity to move from one place to another.
- nomadic
-
generally wanders from place to place, usually within a well-defined range.
- solitary
-
lives alone
- social
-
associates with others of its species; forms social groups.
- colonial
-
used loosely to describe any group of organisms living together or in close proximity to each other - for example nesting shorebirds that live in large colonies. More specifically refers to a group of organisms in which members act as specialized subunits (a continuous, modular society) - as in clonal organisms.
- visual
-
uses sight to communicate
- acoustic
-
uses sound to communicate
- duets
-
to jointly display, usually with sounds in a highly coordinated fashion, at the same time as one other individual of the same species, often a mate
- food
-
A substance that provides both nutrients and energy to a living thing.
- threatened
-
The term is used in the 1994 IUCN Red List of Threatened Animals to refer collectively to species categorized as Endangered (E), Vulnerable (V), Rare (R), Indeterminate (I), or Insufficiently Known (K) and in the 1996 IUCN Red List of Threatened Animals to refer collectively to species categorized as Critically Endangered (CR), Endangered (EN), or Vulnerable (VU).
- carnivore
-
an animal that mainly eats meat
- piscivore
-
an animal that mainly eats fish
- molluscivore
-
eats mollusks, members of Phylum Mollusca
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
References
Ainley, D., T. Telfer, M. Reynolds. 1997. Townsend's and Newell's Shearwaters. The Birds of North America , 297: 1-20.
American Ornithologist's Union, 2009. "Check-list of North American Birds" (On-line). American Ornithologist's Union. Accessed February 03, 2009 at http://www.aou.org/checklist/north/ .
BirdLife International 2008, 2008. "Puffinus auricularis" (On-line). IUCN Redlist of Endangered Species. Accessed February 02, 2009 at http://www.iucnredlist.org/details/144887 .
BirdLife International 2008, 2008. "Puffinus newelli" (On-line). 2008 IUCN Red List of Threatened Species. Accessed February 03, 2009 at http://www.iucnredlist.org/details/144888 .
Brooke, M. 2004. Albatrosses and Petrels across the World . Oxford: Oxford University Press.
Day, R., B. Cooper, T. Telfer. 2003. Decline of Townsend's (Newell's) shearwaters (Puffinus auricularis newelli) on Kauai, Hawaii. The Auk , 120: 669-679.
Day, R., B. Cooper. 1995. Patterns of movement of Dark-rumped Petrels and Newell's shearwaters on Kauai. Condor , 97: 1011-1027.
Dickinson, E. 2003. The Howard and Moore Complete Checklist of the Birds of the World . Princeton: Princeton University Press.
Martinez-Gomez, J., J. Jacobsen. 2004. The conservation status of Townsend's shearwater Puffinus auricularis auricularis. Biological Conservation , 116: 35-47.
Pitman, R., L. Ballance. 2002. The changing status of marine birds breeding at San Benedicto Island, Mexico. Wilson Bulletin , 114: 11-19.
Podolsky, R., D. Ainley, G. Spencer, L. Deforest, N. Nur. 1998. Mortality of Newell's Shearwaters Caused by Collisions with Urban Structures on Kauai. Colonial Waterbirds , 21: 20-34.
Reed, J., J. Sincock, J. Hailman. 1985. Light attraction in endangered procellariiform birds: Reduction by shielding upward radiation. The Auk , 102: 377-383.
Soldaat, E. 2007. "Puffinus shearwaters" (On-line). Seabird Osteology Pages. Accessed February 03, 2009 at http://www.shearwater.nl/seabird-osteology/General%20chapters/ssg_introduction.htm .
Spear, L., D. Ainley, N. Nur, S. Howell. 1995. Population size and factors affecting at-sea distributions of four endangered procellariids in the tropical Pacific. Condor , 97: 613-638.