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Additional
Photos
branchia
oral tentacles
anal papilla
juvenile
early transitional
late transitional
flared mantle
with opaque flecks
dark
senescent?
swimming
with shrimp
predation
mating
egg mass
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GALLERY
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Hexabranchus sandwichensis (Gray, 1850)
Maximum size: 300 mm.
Identification:
This
species
is
the largest and, perhaps, most widely recognized
Hawaiian nudibranch. Mature animals are crimson
mottled with pink and sometimes flecked with opaque white. When the
animal is at rest, the mantle
edge is rolled but, when disturbed, it's flared revealing a
dark red margin. The oral tentacles are unusual in that they
resemble tiny hands. As with other species of Hexabranchus, it undergoes complex
changes in color and form as it matures. Very young animals are cream,
decorated with violet spots, and lack broad lateral extensions of the
mantle. Maturing animals gradually lose the violet spots and develop
cream mottling. The background darkens to golden yellow-orange, then
crimson. A "frosting" of opaque white flecks may appear on the notum in very old animals.
Rarely, mature animals may be yellowish. (Note 1) It can be distinguished from Hexabranchus
aureomarginatus by the lack of a yellow marginal band at all
sizes. See the article
on this site for further discussion.
Natural history: Hexabranchus sandwichensis is a common
nocturnal species that shelters under rocks or in crevices during the
day.
It lives in protected to moderately exposed rocky areas from < 1 to
55
m (< 3 to 180
ft) and has also been photographed
from HURL submersibles at 63
m (207 ft). Rarely, it may be seen in the open during the day. (Note 2) Like many dorids, it obtains protective chemicals from its
sponge food and Kay (1979) reports that it feeds on the calcareous sponge Leucetta solida. (Note 3)
In
addition,
it
is
one of the few that can leave the sea floor and swim
when threatened--the "dance" referred to in its common name. When
swimming, the mantle
margin is unrolled to reveal a dark red band and there is strenuous
dorso-ventral flexing, perhaps serving to elicit a startle response or
advertise its toxic nature to potential predators. The commensal
imperial shrimp, Zenopontonia rex (= Periclimenes imperator), is
sometimes found living on its body, often among the gills. (see photo) Its egg mass
is large and conspicuous consisting of 1-5 bright pink coils often laid
in an exposed location such as the top of a piece of coral rubble or
rocky spur (see
photo). The eggs are very small and deposited in discrete packets of
a few dozen that are embedded in the ribbon. Laying may occur primarily
at night. As the eggs develop, the egg mass fades to a pale pink and
may become worn and tattered-looking. Defensive
chemicals are deposited in the egg mass at much higher concentrations
than in the nudibranch itself (Scheuer, 1990) so the masses are
protected
during their
week-long development. Most predators are deterred by the chemicals (Note 4) but Favorinus japonicus, a species
of aeolid nudibranch, is not. (Note 5)
Distribution:
Big Island, Maui, Kahoolawe, Lanai, Oahu, Kauai, Niihau, French Frigate Shoals and Midway (also Johnston Atoll).
Taxonomic notes:
This
species
is
listed as Hexabranchus
sanguineus (Rupell and Leuckart, 1831) in Kay, 1979 and Kay & Schoenberg-Dole, 1991.
It's listed as both Hexabranchus marginatus and Hexabranchus cf. pulchellus in Kay & Young, 1969. Animals referred to as H. pulchellus in Kay, 1979 and illustrated
as such in Bertsch and Johnson, 1981 are juveniles. It is also listed as H. sanguineus in Hoover, 1998 &
2006 (right photo) (corrected in 5th printing). The drawing labelled Hexabranchus sp. in Edmondson, 1946 is probably this species. Doris cardinalis, Doris sandwichiensis, Hexabranchus cardinalis and Hexabranchus tinkeri are also synonyms. It is one of the
species commonly referred to as
the "Spanish dancer" in Hawaii and elsewhere. In the 5th printing,
Hoover, 2006 refers to is as the "redmargin Spanish dancer." It was previously listed on this site as Hexabranchus pulchellus. Various other sources for Hawaii typically list it as either H. sanguineus or H. pulchellus. See Tibirica, et. al (2023) for further discussion.
Photo: PF:
about 250 mm: Molokini Islet, Maui; Nov. 20, 2008.
Observations and comments:
Note 1: Kauai Anew includes photos of an unusual yellowish animal. Except for the atypical background color, its characteristics match this species.
Note 2:
On Maui, we've only seen this species resting in the open by day on one
or two occasions. However, in May, 2012 several animals were observed
crawling in the open by day in tide pools and the shallow subtidal on
Kauai. This suggests more flexible behavior than our Maui observations
with at least some diurnal (or crepuscular?) activity. The daytime
sightings were in the morning under cloudy conditions.
Note 3: In spite of its chemical
defense, we have seen it preyed upon by sponge crabs (Tumidodromia dormia), the splendid pebble
crab (Etisus splendidus), the banded spiny lobster (Panulirus
marginatus) and the red swimming crab (Gonioinfradens paucidentata): all nocturnal hunters. Regarding the last, PF reported: "...Gonioinfradens paucidentata
gripping a live Spanish Dancer which was trying to get away. You can
see some damage to the nudibranch near the right claw. After awhile the
crab got spooked and ran off with the nudibranch in its claws." (see
photo)
Note 4: Fish
that have not learned of their toxicity may bite the egg masses but
almost
always let go without completely removing a piece of the ribbon, leaving
it dangling. We
have also seen green sea turtles (Chelonia
mydas) mouthing egg masses but not eating them.
Note 5:
John Horn photographed an egg mass on Kauai that had been colonized by
micro-algae. There's some uncertainty in assigning the mass to H. sandwichensis (John reports that H. aureomarginatus
is more common in the area). However, the height of the ribbon is more
typical of this species. As far as I can recall, we've never seen micro-algae on an egg mass.
Perhaps, some factor could have delayed hatching providing sufficient
time for it to colonize? Alternatively, if it's actually an H. aureomarginatus mass, perhaps the masses of that species take longer to hatch or are more vulnerable to algal growth than H. sandwichensis
masses? (see photos)
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