ArticlePDF Available

Intertidal records of ‘sea slugs’ (nudibranchs and allied opisthobranch gastropods) from northern North Island, New Zealand

Authors:
  • Geomarine Research

Abstract and Figures

We record the intertidal or shallow subtidal (<2–3 m) occurrence of 78 species of ‘sea slug’ from northern North Island of New Zealand. One (Goniodoris n.sp.) is recorded for the first time. The majority are briefly described, illustrated and their records plotted on maps. Three taxa are probably undescribed new species. All except two (Aphelodoris sp., Trinchesia reflexa) of the species have been recorded from along the east coast (warmer water Aupourian Province) but only 44% (34 spp.) have been recorded from the west coast (cooler Cookian Province). The highest diversity of intertidal/shallow water ‘sea slugs’ has been recorded from the Leigh area (47 spp.), Bay of Islands (42 spp.) and Great Barrier Island (38 spp.) reflecting the intensity of survey (Leigh) and diversity of habitats on the warmer coast (latter two). Of the harbours, 33 spp. are recorded from the Waitemata, 27 spp. from Parengarenga and 24 spp. from the Manukau, reflecting the intensity of survey (Waitemata, Manukau) and unusual warm conditions of far north Parengarenga. Of the ‘sea slugs’ recorded herein, 45% are endemic to New Zealand and nine of these are endemic to northern New Zealand.
Content may be subject to copyright.
Records of the Auckland Museum 50: 2015
INTRODUCTION
The common name ‘sea slug’ conjures up repulsive
images of dull, slimy creatures, but in reality, while some
are cryptic, many display brilliant colours and dramatic
colour combinations more like butteries. Readers are
strongly recommended to complement the black and white
images presented here with the coloured images in Willan
et al. (2010) or online sites (http://www.seaslugforum.
net/, http://www.nudipixel.net/, http://www.ianskipworth.
com/suig/nudis.html). Detailed descriptions of anatomy,
feeding and reproduction are given in Willan and Morton
(1984) and Willan et al. (2010). Enderby and Enderby
(2005) presented an illustrated account of endemic
nudibranchs in New Zealand.
Sea slugs seen by scuba divers at subtidal depths down
to ~30 m have been well reported (Willan and Morton
1984; Willan et al. 2010; http://www.seaslugforum.net/),
but intertidal and shallow subtidal species from snorkel
depths (0–3 m) have not been documented so fully. Those
sea slug species that have been recorded from northern
New Zealand, but only from subtidal scuba depths (>
3 m) have not been included here. Abundances of some
species appear to dier between intertidal and subtidal
populations, e.g. Ceratosoma amoenum and aeolids are
more common subtidally than intertidally (pers. obs.),
so this compilation of our observations over many years
compliments what is already well known and helps
advance our knowledge of the geographic distribution,
ecology and in some instances the life history of sea
slugs in northern New Zealand.
What is meant by a ‘sea slug’
The name ‘sea slug’ is usually applied to a number
of marine gastropods that are shell-less or outwardly
appear to have no shell. It seems that reduction in size
or loss of a shell has occurred on a number of occasions
in dierent evolutionary lineages of opisthobranch
gastropods. Here we adopt a pragmatic approach for
our more general readers and use the term ‘sea slug’ for
slug-like marine opisthobranch gastropods that have no
shell, or their shell is internal or where their shell is too
small for the soft parts to be retracted into it. The largest
group numerically is the Nudibranchia, where there
is no shell as an adult and the often brightly-coloured
animal has external gills. Some of the sea slugs included
here (Family Aplysiidae) are also known as sea hares
because of their apparent resemblance to a sitting hare.
Abstract
We record the intertidal or shallow subtidal (<2–3 m) occurrence of 78 species of ‘sea slug’ from
northern North Island of New Zealand. One (Goniodoris n.sp.) is recorded for the rst time. The
majority are briey described, illustrated and their records plotted on maps. Three taxa are probably
undescribed new species. All except two (Aphelodoris sp., Trinchesia reexa) of the species have
been recorded from along the east coast (warmer water Aupourian Province) but only 44% (34
spp.) have been recorded from the west coast (cooler Cookian Province). The highest diversity
of intertidal/shallow water ‘sea slugs’ has been recorded from the Leigh area (47 spp.), Bay of
Islands (42 spp.) and Great Barrier Island (38 spp.) reecting the intensity of survey (Leigh) and
diversity of habitats on the warmer coast (latter two). Of the harbours, 33 spp. are recorded from
the Waitemata, 27 spp. from Parengarenga and 24 spp. from the Manukau, reecting the intensity
of survey (Waitemata, Manukau) and unusual warm conditions of far north Parengarenga. Of the
‘sea slugs’ recorded herein, 45% are endemic to New Zealand and nine of these are endemic to
northern New Zealand.
Keywords
New Zealand; Hauraki Gulf; Northland; nudibranchs; ‘sea slugs’; biogeography.
Intertidal records of ‘sea slugs’ (nudibranchs and
allied opisthobranch gastropods) from northern
North Island, New Zealand
Margaret S. Morley Auckland War Memorial Museum
Bruce W. Hayward Geomarine Research (New Zealand)
51–93
52
They usually have a long neck bearing a head with a pair
of enrolled rhinophores and enrolled tentacles on each
side of the mouth. In addition there is one other common
intertidal slug-like gastropod (Scutus) in northern New
Zealand that is not an opisthobranch but an unrelated
ssurellid which we have included.
Survey area, methods and sources of records
The area covered by this study is northern New Zealand
and extends northwards from the Manukau Harbour,
Firth of Thames and northern Coromandel Peninsula
and includes Great Barrier Island, but not the Three
Kings or Kermadec Islands (Fig. 1). The majority of
the records reported here have been made by MSM and
BWH since 1995 while undertaking semi-quantitative
assessments of the intertidal biota of dierent parts of
the shoreline of the region (Appendix 1). The surveys
were not specically targeted for nudibranchs. Each
record placed on the maps (Figs 1–24) relates to one or
more ndings within a coastal stretch of 0.5–2 km that
was typically surveyed over a 2–4 hr period either side
of a spring low tide. This paper also includes records
from major study areas where the total biota has already
been documented and published. These are, from north
to south: Parengarenga Harbour (Hayward et al. 2001),
Ahipara (Hayward et al. 2004), Kawerua (Hayward et
al. 1995), Bay of Islands (Morley and Hayward 1999),
Whananaki (Hayward et al. 2012), Great Barrier Island
(Morley and Hayward 2009), Whangapoua, Coromandel
Peninsula (Hayward et al. 2014), Waitemata Harbour
(Hayward et al. 1999), Tamaki Estuary (Hayward and
Morley 2008), Waitakere Ranges and northern Manukau
Harbour (Hayward and Morley 2004). Many additional
sites in the southern half of the region have also been
Margaret S. Morley & Bruce W. Hayward
Former name Present name
Aeolidiella indica (Bergh, 1888) Anteaeolidiella lurana
Aplysia dactylomela Rang, 1828 Aplysia argus
Archidoris wellingtonense Abraham, 1877 Doris wellingtonensis
Bouvieria ornata (Cheeseman, 1878) Berthella ornata
Bulla angasi Bulla vernicosa
Bursatella glauca (Cheeseman, 1878) Bursatella leachii
Chromodoris amoena Cheeseman, 1886 Ceratosoma amoenum
Chromodoris aureomarginata Cheeseman, 1881 Goniobranchus aureomarginatus
Ctenodoris abellifera (Cheeseman, 1881) Doris granulosa
Cuthona beta (Baba & Abe, 1964) Trinchesia beta
Cuthonia reexa Miller, 1977 Trinchesia reexa
Dendrodoris denisoni (Angas, 1864) Dendrodoris krusensternii
Dendrodoris gemmacea (Alder & Hancock, 1864) Dendrodoris krusensternii
Dermatobranchus pulcherrimus Miller & Willan, 1986 Dermatobranchus rubidus
Doriopsis abellifera (Cheeseman, 1881) Doris granulosa
Glossodoris amoena (Cheeseman, 1886) Ceratosoma amoena
Glossodoris aureomarginata (Cheeseman, 1881) Goniobranchus aureomarginatus
Hydatina albocincta (van der Hoeven, 1839) Hydatina zonata
Okadaia cinnabarea (Ralph, 1944) Vayssierea cinnabarea
Placida aoteana (Powell, 1937) Placida dendritica
Pleurobranchaea novaezealandiae Cheeseman, 1878 Pleurobranchaea maculata
Pupa kirki (Hutton, 1873) Pupa affi nis
Rostanga rubicunda (Cheeseman, 1881) Rostanga muscula
Scutus antipodes Montfort, 1810 Scutus breviculus
Spurilla drusilla (Bergh, 1900) Aeolidia drusilla
Trippa molesta Miller, 1989 Atagema molesta
Umbraculum sinicum (Gmelin, 1791) Umbraculum umbraculum
Table 1. Published name combinations used for some intertidal New Zealand sea slugs that have been
changed since 1980. For earlier synonymies refer to Powell (1979).
53Intertidal records of ‘sea slugs’
surveyed by the authors but the results have not yet been
published (Appendix 1).
Where the identication of a specimen was not
immediately obvious, it was placed in sea water to
encourage extension and movement. This allowed
observation of its colour, shape, gill, cerata, rhinophores
and tentacles. These characteristics were recorded
and sometimes the specimen was photographed. A
few unfamiliar specimens were collected as vouchers,
narcotised, preserved and placed in the wet collections
of the Marine Department of the Auckland War Memorial
Museum (specimens prexed by MA in Appendix 1).
Historical locality records dating back to 1980
made by one of us (MSM, Appendix 1) have also been
included, and include observations of species made
during snorkelling at shallow subtidal depths down to
1–3 m. These unpublished records consist of drawings,
photographs and notes. A number of additional locality
records have been added from the collection database of
the Marine Department, Auckland Museum and Te Papa
Museums’ web-based database as detailed in Appendix 1.
Drawings
Most of the recorded species are accompanied by a
drawing usually by MSM. These are generally based on
actual live specimens that were placed in sea water and
sketched. In some instances these sketches have then
been augmented by observations made on live specimen
photographs on the internet. A few species were not
available for drawing from either our specimens or
notes, and these have been redrawn from Willan and
Morton (1984). Where there is no locality given in
the caption to a drawing, the specimen drawn was not
found live by us, and has been drawn from a photograph
from elsewhere.
Previous work
Most of the more common larger sea slugs seen
intertidally around northern New Zealand were described
in the 19th century. Of particular interest are six that were
described by the then Director of Auckland Museum,
Thomas Cheeseman between 1878 and 1886. Because
it was impossible to preserve their colour, he had his
sister Emma Cheeseman draw iconotypes of some of
these in 1881-2, which are held by the Museum (Powell
1941). In 1979, Auckland Museum’s Arthur ‘Baden’
Powell recorded a total of 73 species of ‘sea slugs’ from
New Zealand, but some of these have subsequently
been synonomised. Since then Univeristy of Auckland
professor Michael Miller and his graduate students, Derek
Challis, Bill Rudman and Richard Willan, have published
over twenty papers on the nudibranchs of New Zealand,
a number of them described as new. In 1984, Richard
Willan and John Morton provided general descriptions
and illustrated 150 species of ‘sea slugs’ from New
Zealand. Updated information and colour photographs of
87 New Zealand ‘sea slugs’ were provided in Willan et
al. (2010). With molecular studies the taxonomy of sea
slugs continues to change and we have tried to use the
latest synonymies (Table 1).
SEA SLUGS RECORDED INTERTIDALLY FROM
NORTHERN NEW ZEALAND
Classication follows Spencer et al. (2014) and the World
Register of Marine Species (http://www.marinespecies.
org/). The species recorded below are summarised in
Appendix 2. The known global distribution of species is
mostly taken from Willan and Morton (1984) and Willan
et al. (2010).
Opisthobranchia
Acteonidae
Pupa anis (A. Adams, 1855) Fig. 1
Average animal length 15 mm. The shell is elongate-
conic, with narrow punctate spiral grooves cutting the
surface into uneven flat-topped ribs. The interstices
are crossed by fine axial threads and a double fold is
prominent at the base of the columella (Powell 1979,
p 269). The animal is translucent and creamy white.
This species is rare intertidally, where it lives buried
in low tidal sediment. It is more common in dredged
material below low tide to depths of 300 m. In our
records we have found it on semi-protected and open
sandy coasts along the full length of eastern Northland
but only off Ahipara (dredged at 40 m) on the west
side, although it has been collected by others from off
Awhitu Peninsula. Its recorded range extends south to
Raglan on the west coast and down to East Cape on the
east coast (Te Papa Museum records). It is widespread
in the southern hemisphere.
Bullinidae
Bullina lineata (Gray, 1825) Fig. 1
Average animal length 20 mm. The fragile shell is white
with delicate red axial and spiral lines and a sculpture
of incised spirals. The luminous animal is blue, with its
mantle and head shield exquisitely margined by iridescent
greenish-blue. It is thought to prey on polychaete worms
(Rudman 1998). A pair of live specimens was observed
laying spawn at Paua, Parengarenga Harbour at a depth
of 2 m. B. lineata is rare intertidally in New Zealand,
but common in Australia and the Indo-Pacic on ne
sandy substrates down to depths of 10 m. Here we
record it in sporadic occurrences down the full length of
the east coast of Northland and on Great Barrier Island.
Its southernmost record is the Aldermen Islands (Te
Papa Museum records). Dead shells have washed in at
East Cape.
Aplustridae
Hydatina physis (Linnaeus, 1758) Fig. 1
Average animal length 75 mm. Its common name
rose-petal bubble shell is well chosen; the pink animal
is much larger than its fragile shell and its intense blue
margin is exquisitely gathered into numerous rues.
The ground colour of the globose shell is bu, with thin
brown spiral lines and a sunken spire. In August 1986, a
54 Margaret S. Morley & Bruce W. Hayward
Figure 1. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Pupa affin
(Great Barrier, Tryphena), Bullina lineata (Parengarenga, Paua), Hydatina zonata (Northland, Rangiputa) and Hydatina
physis (Parengarenga, Paua). Scale bars 10 mm long.
pair was observed mating in channel mud in a depth of
2 m at Paua, Parengarenga Harbour (MSM pers. obs.).
In August 2000 over 100 specimens of a range of sizes
were found alive over a two day period. Some of these
were oating, some crawling on the mud. The animals all
had light lemon coloured bodies with a white border to
the mantle (Raven 2000). This species lives intertidally
down to 10 m in semi-sheltered localities and only rarely
washed ashore. We have records of dead shells washed in
at four localities along the northeast coast of Northland –
Parengarenga, Rangiputa, Bay of Islands and Tutukaka.
It is widespread throughout the tropical and subtropical
Indo-Pacic and Atlantic Oceans.
Hydatina zonata (Lightfoot, 1786) Fig. 1
Average animal length c. 35 mm. The fragile inated
shell is very variable in coloration. Some have four
dark brown radial stripes on a white ground (Powell,
1979), while others have various combinations of
narrow brown and and white spirals (Rudman 2004).
A single empty shell of this species was recorded from
Tutukaka in April 1970 by Powell (1979, p. 270). For 3
to 4 years from 2003-2006, extremely rare dead shells
were washed in at Rangiputa and Rangaunu Harbour in
the Far North (Peter Poortman pers. comm.). The only
recorded live specimen in New Zealand was found at
Paua, Parengarenga Harbour in 2000 (Raven 2000). The
animal was pinkish mauve with a white border. There
are no recent records of this species in New Zealand. It
is also known from Japan and northern Australia.
55Intertidal records of ‘sea slugs’
Figure 2. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Cylichna
thetidis (Great Barrier, Whangaparapara), Philine angasi (Manukau Harbour, Cornwallis), Philine auriformis (Tamaki
Estuary, Farm Cove) and Retusa oruaensis (Northland, Bland Bay). Scale bars 10 mm long unless stated otherwise.
Cephalaspidea
Cylichnidae
Cylichna thetidis Hedley, 1903 Fig. 2
Average animal length 10 mm. The shell is small, white
with sub-parallel sides and a low spire. There are usually
ne spiral lines present. It lives in sand intertidally down
to 55 m (Willan and Morton 1984). We have found it
in low numbers in our surveys in shell sand throughout
the east coast of Northland and Auckland, but not in the
inner gulf. We have occasionally sieved it alive in low
tide sediment. It is also known from further south around
the North Island and in Australia.
Philinidae
Philine angasi (Crosse and Fischer, 1865) Fig. 2
Average animal length 50 mm. The animal is a rm,
white, carnivorous slug that buries in soft sediment at or
below low tide down to 7 m (Willan and Morton 1984).
The internal shell is smooth, thin and fragile with a large
body whorl. We have found specimens buried in extreme
low tidal sandy mud at Cornwallis and Wattle Bay, in the
Manukau Harbour. We have scattered records from right
around Northland and Auckland with its range extending
south to East Cape. It also occurs in Australia.
Philine auriformis Suter, 1909 Fig. 2
Average animal length 25 mm. This smaller relative
of P. angasi is distinguished from it by delicate spiral
56
sculpture on the internal shell. This can be seen under
the microscope through the tissues of the living animal.
It lives buried in sediment at low tide and deeper. We
have scattered records from right around Northland and
Auckland. It occurs right around the three largest islands
of New Zealand. It is an invader on the west coast of
United States of America (Krug et al. 2012).
Retusidae
Retusa oruaensis (Webster, 1908) Fig. 2
Average animal length 3 mm. The tiny, white cylindrical
shell is parallel-sided, and the animal creamy white.
This endemic species lives buried in sediment usually
below low tide and down to 200 m (Willan and Morton
1984). At Bland Bay, Northland it was live in sediment
in a water depth of 1 m at low tide. We have records of
washed up dead shells from right around northern New
Zealand and its range extends around the South Island
and out to the Chatham Islands. Type locality is Orua
Bay, Manukau Harbour.
Aglajidae
Melanochlamys cylindrica Cheeseman, 1881 Fig. 3
Average animal length 25 mm. This black cylindrical
slug looks like a twig, making it easily overlooked as
it crawls on coralline turf. The parapodia are closely
wrapped round the body. It preys on nereid worms
(Willan and Morton 1984). There is a small, vestigial,
internal shell. This endemic slug is often found in low
numbers during our searches in mid to low intertidal
Margaret S. Morley & Bruce W. Hayward
Figure 3. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Melanochlamys
cylindrica (Whangaparaoa, Fishermans Rock), Melanochlamys lorrainae (Parengarenga, Paua), Philinopsis taronga
(Tamaki Estuary, Farm Cove), Runcina katipoides (Northland, Pataua), Pseudoilbia lineata and Runcinella zelandica. Scale
bars 10 mm long unless stated otherwise.
57Intertidal records of ‘sea slugs’
pools both on muddy shore platforms and on the open
coast. We have scattered records along the east coast of
Northland and Auckland and in Manukau Harbour, with
its most common occurrence on the west coast of the
Hauraki Gulf. It occurs right around both main islands
of New Zealand.
Melanochlamys lorrainae (Rudman, 1968) Fig. 3
Average animal length 25 mm. Unlike M. cylindrica, M.
lorrainae is rare. The animal is white to grey with yellowish
tinges and has only been recorded in soft sediment. The type
locality is from dredged sediment o Wattle Bay, Awhitu
Peninsula, Manukau Harbour. A specimen, observed by
MSM at Paua, Parengarenga Harbour, emerged covered
in mucus from the low tidal muddy sand on the turning
tide. Molecular phylogenic analysis on another specimen
collected from low tide sand at Wattle Bay showed that
this is a distinct species from M. cylindrica (Krug et al.
2008). The endemic M. lorrainae has been recorded in
harbour sediment from scattered localities down the east
side of northern New Zealand and in Manukau Harbour.
Its southern most record is in the Marlborough Sounds (Te
Papa Museum database).
Philinopsis taronga Allan, 1933 Fig. 3
Average animal length 30 mm. This uncommon aglajid
is brown with an irregular, black, reticulated pattern
enclosing yellow spots. The parapodia and head shield
are edged with yellow to orange. The body is often folded
centrally resulting in a semi-curled position. There is
an internal shell. It is found living in low tide mud or
sand on sheltered beaches A possible prey species is
Haminoea zelandiae which is common in the same area
of the Tamaki Estuary. We have sporadic records from
the east coast of northern New Zealand. It is also known
from Australia.
Runcinidae
Pseudoilbia lineata Miller and Rudman, 1968 Fig. 3
Average animal length 3 mm. The front and back of the
animal are pale white and the remainder has ne black
longitudinal bands. There is a narrow orange-red bar
behind these bands. It lacks a gill, shell and gizzard
plates. The holotype was found in an intertidal pool
on coralline turf at Langs Beach, Bream Tail, in 1963
(Miller and Rudman 1968). No further specimens have
been found.
Runcina katipoides (Miller and Rudman, 1968) Fig. 3
Average animal length 5 mm. This tiny elongate nudibranch
is seldom seen unless its habitat in coralline turf is carefully
examined. It is a herbivore (Sea Slug Forum). The digestive
gland shows as a bright red, central stripe on the dorsum
of the dark grey body. At the rear of the notum a white
patch denotes the internal shell. The anterior area around
the eyes is clear. The margin of the body has a narrow blue
phosphorescent border (MSM pers. obs.). This species is
a direct developer. Spawn is laid, with the black centred
eggs in a ball of clear jelly with a diameter of 2 mm (MSM
pers. obs.). It is found on hard substrates from protected
harbours to open coastal localities intertidally down to 14
m depth (Willan and Morton 1984). This endemic species
is only known from the east coast of northern Northland
between the Bay of Islands and Opoutere on the east coast
of Cormandel Peninsula. Its type locality is Taurikura Bay,
Whangarei Harbour.
Runcinella zelandica Odhner, 1924
Average animal length 5 mm.The animal has a dark green
background with irregular salmon pink longitudinal
bands and sometimes irregular white patches. Corners
of the head are folded into tentacular processes that give
it a bilobed appearance. It has large gills that project
out from under the posterior edge of the mantle. This
endemic species is most commonly seen feeding on the
green alga Codium dichotomum in the sublittoral fringe
down to about 3 m depth (Willan 1981). It was described
from Cape Brett (Bay of Islands) but is most commonly
seen at the Poor Knights Islands.
Bullidae
Bulla quoyii Gray, 1843 Fig. 4
Average animal length 50 mm. The distinguishing
features of this brown shell are the spiral grooves at
the anterior end. It is common and widespread in our
study area on sheltered beaches buried in muddy pockets
among coralline algae or in seagrass. Empty, brown
shells are often washed up on beaches, but the animal
is seen less often since it is nocturnal. It occurs right
around northern New Zealand, being most abundant on
the more sheltered harbour shores. It is also known from
southern Australia.
Bulla vernicosa Gould, 1859 Fig. 4
Average animal length 30 mm. The shell is more solid
than B. quoyii and has a glossy surface. There are no
spiral striations around the base of the shell and the outer
lip is attened to slightly concave rather than convex.
The dark speckles often form two or three spirals bands
on the shell (Willan et al. 2010). It lives subtidally at
2–10 m depth on semi-exposed soft shores with rare
empty shells washed ashore on Northland’s east coast.
It also occurs at the Three Kings and Kermadec Islands
and elsewhere in the subtropical Pacic.
Haminoeidae
Haminoea zelandiae (Gray, 1843) Fig. 5
Average animal length 30 mm. The dark grey animal
shows through the thin, fragile, white shell, so living
specimens often go unnoticed. This intertidal grazer is
common on protected harbour shores. It lives in muddy
patches between coralline turf and also in eel grass
Zostera, feeding among lamentous algae, probably
on their epiphytic diatoms (Willan and Morton 1984).
This endemic species is widespread on sheltered coasts
right around northern New Zealand as far south as the
northern South Island.
58
Aplysiomorpha
Aplysiidae
Aplysia argus (Ruppell and Leuckart, 1830) Fig. 6
Average animal length 300 mm, internal shell 70 mm.
This large handsome sea hare is olive green with a
network of black reticulating lines. Its distinguishing
feature is the black patch on the tail. The erect parapodia
are connected low down posteriorly (Willan et al. 2010).
It lives predominantly intertidally on open rocky coasts
where it grazes on red and green algae (Willan 1979a). The
animal secretes purple ink if disturbed. We have sporadic
records of single specimens on the east coast of northern
New Zealand and from Ahipara on the west coast. It is
cosmopolitan in warm temperate and tropical seas.
Aplysia juliana (Quoy and Gaimard, 1832) Fig. 6
Average animal length to 250 mm, internal shell 30 mm.
This uncommon sea hare varies in colour from chocolate
brown to black with a scattering of white spots on the
external surface of the parapodia, head and tail. It crawls
like an inch worm and lives from low tide to 3 m. We
have tidal pool records of several specimens at Sentinel
Rock, Mangawhai Heads and several snorkelling records
also from the east coast of northern New Zealand as far
south as Wellington (Te Papa Museum database). It is
cosmopolitan in warm temperate and tropical seas.
Aplysia keraudreni Rang, 1828 Fig. 6
Average animal length 250 mm. This large sea hare is
kelp-brown with white streaks and mottles overlaid by
a black netted pattern. There is a small internal shell.
Margaret S. Morley & Bruce W. Hayward
Figure 4. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Bulla quoyi
(Waitemata Harbour, Musick Pt) and Bulla vernicosa (Northland, Spirits Bay). Scale bars 10 mm long.
59Intertidal records of ‘sea slugs’
Figure 5. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Haminoea
zelandiae (Tamaki Estuary, Farm Cove). Scale bars 10 mm long.
Its habitat is on open rocky coasts and harbour mouths
from intertidal down to 18 m depth. It feeds on red algae.
We have scattered intertidal records from the east coast
of northern New Zealand, but it also occurs in central
and southern parts of the country. Its range is the south-
western Pacic, including Australia.
Aplysia parvula Guilding, 1863 Fig. 6
Average animal length 60 mm. This is the smallest
sea hare in New Zealand. It is kelp-brown dotted with
white, and the parapodia, rhinophores and oral tentacles
are edged with a narrow black line. The parapodia are
continuous forming a cup, only open in the front. It has
the largest eye spots of all the Aplysia species. All our
records are from the east coast of northern New Zealand
but it is known from right around the two main islands.
It is cosmopolitan in warm temperate and tropical seas.
Bursatella leachii Blainville, 1817 Fig. 7
Average animal length 80 mm. This common sea hare
is camouaged by feather-like processes. The body is
brownish-green, decorated with black speckles and vivid
emerald green patches. It can be found in high numbers
in protected situations such as harbours, estuaries and
on Zostera beds, especially in summer when they come
inshore to breed (Willan and Morton 1984). All our
records are from the east coast of northern New Zealand,
particularly around the inner Hauraki Gulf. In 2000 there
was a population explosion at Howick Beach, Waitemata
Harbour following a major algal bloom (Morley et al.
2001 p.16). Its southernmost New Zealand record is
Cape Kidnappers, Hawkes Bay (Te Papa Museum
database). Its range includes tropical and subtropical
waters of the South Pacic.
60
Figure 6. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Aplysia
argus (Waiheke Is, W Bay), Aplysia juliana (Northland, Mangawhai Heads), Aplysia keraudreni (after Willan and Morton,
1984, p. 32) and Aplysia parvula (Northland, Bland Bay). Scale bars 10 mm long unless stated otherwise.
Margaret S. Morley & Bruce W. Hayward
61Intertidal records of ‘sea slugs’
Figure 7. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Bursatella
leachii (Tamaki Estuary, Bucklands Beach), Dolabrifera brazieri (Northland, Taupo Bay) and Stylocheilus longicauda (after
Willan and Morton, 1984). Scale bars 10 mm long unless stated otherwise.
Dolabrifera brazieri G.B. Sowerby II, 1870 Fig. 7
Average animal length 80 mm, shell 8 mm. A
distinguishing feature of this sea hare is its attened
shape and wide foot allowing it to closely conform to the
rock, often in intertidal pools. Its green body is marbled
with brown, white and pink on low rounded papilli
(Willan and Morton 1984). It has only been found on
the open eastern coast of northern New Zealand. This
tropical species arrived in New Zealand in the 1970s
(Willan and Morton 1984, p 37)).
Stylocheilus longicauda (Quoy and Gaimard, 1825) Fig. 7
Average animal length 60 mm. This is a rare sea hare
with an elongate body and a long narrow tail. It can
occur with Bursatella leachii and has similar dendritic
processes. The green body has ne longitudinal brown
lines and spots that are ringed with black. It lives
intertidally down to 30 m, on protected to semi-exposed
coasts where it feeds on lamentous green and blue green
algae. It occurs rarely along Northland and Auckland’s
east coast (Willan and Morton 1984). It is cosmopolitan
in warm temperate and tropical seas.
Sacoglossa
Sacoglossans are suctorial herbivores on green algae
(Powell 1979, p. 283). Most saccoglossans are green,
the pigment being derived from their food. The radula
is adapted into a single line of teeth ideal for piercing
the algal cells.
62
Figure 8. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Ercolania
felina (Waitakere coast, Powell Bay), Steliger smaragdinus (redrawn from Sea Slug Forum web site), Placida dendritica
(Waitemata Harbour, Mangemangeroa Estuary) and Elysia maoria (Kauri Point, Waitemata Harbour) and Elysia n.sp.
Scale bars 10 mm long unless stated otherwise.
Limapontiidae
Ercolania felina (Hutton, 1882) Fig. 8
Average animal length 10 mm. This small sacoglossan
is grey to black with a central white strip on the dorsum
and around the eyes. There are two rows of club-shaped
appendages similar to cerata down each side of the body.
Grooves are present on the underside of the head, body
and rhinophores to secure it to the bright green strands of
the green alga Chaetomorpha aerea on which it feeds and
lays its spawn coils (Morley 2004). It lives in high tidal
rock pools on exposed coasts. In northern New Zealand
this endemic species has been recorded from both coasts
of Auckland, common at times on the Waitakere coast. It
occurs right around both main islands of New Zealand.
Stiliger smaragdinus Baba, 1949 Fig. 8
Average animal length 15 mm. A seldom seen
sacoglossan because it is almost invisible on its food
source. Its bright green bulbous appendages are a
perfect match for the alga Caulerpa geminata. The
rhinophores have white tips. The algal pigments are
retained in the digestive system and help colour the
animal. In New Zealand it has only been recorded
from Goat Island, Leigh. It is also found in Japan and
Australia (Sea Slug Forum).
Placida dendritica (Alder and Hancock, 1843) Fig. 8
Average animal length 10 mm. This small sacoglossan
is a suctorial feeder on green alga, most commonly
on Codium convolutum. It has a attened green body
Margaret S. Morley & Bruce W. Hayward
63Intertidal records of ‘sea slugs’
with numerous slender cerata. It lives on rocky reefs
in semi-sheltered localities, from low intertidal to 14
m depth (Willan et al. 2010). This species occurs right
around New Zealand. It is cosmopolitan in warm to cool
temperate seas.
Placobranchidae
Elysia maoria Powell, 1937 Fig. 8
Average animal length 16 mm. This sacoglossan is dark
green with parapodia that meet in the midline. Under the
microscope a detailed pattern of red and white spots is
visible. It is found, intertidally to 4 m, on rocky reefs
in semi-sheltered localities where it lives on and under
Codium convolutum and C. fragile ssp. tomentosoides.
The animal blends so well among the green alga it can
only be detected by the presence of its white spiral
spawn coil. In our studies we have found it around the
sheltered shores of Hauraki Gulf, being most common in
the Waitemata Harbour, especially at Bucklands Beach,
Tamaki Estuary. Its range includes Australia.
Elysia n.sp. Fig. 8
This is an undescribed species which diers from E.
maoria by its colouration of pale green parapodia
with blue spots and a brilliant blue-green margin. It is
recorded from oshore islands, east of Northland (Fig.
8) by Willan et al. (2010; pers. comm.). Also recorded
from Australia and Norfolk Island.
Figure 9. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Berthella
medietas (Waitemata Harbour, Achilles Pt), Berthella ornata (Waitemata Harbour, Musick Pt) and Berthellina citrina
(Coromandel Peninsula, Fletchers Bay). Scale bars 10 mm long.
64
Figure 10. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of
Pleurobranchaea maculata (Motuihe Island). Scale bars 10 mm long.
Pleurobranchomorpha
Pleurobranchidae
Characteristics of pleurobranchs are their triangular oral
veil, rolled rhinophores, either arising close together or
separated, and a gill on the right side.
Berthella medietas Burn, 1962 Fig. 9
Average animal length 25 mm. It has a small internal
shell. This orange to bronze pleurobranch has a
characteristic porous texture which distinguishes it
from Berthellina citrina. Under the microscope glassy
spicules can be seen interspersed between the pores.
The rhinophores are conjoined above a wide oral veil. It
lives on partly sheltered and open rocky coasts, down to
depths of 5 m. We have found it in low numbers under
low tidal boulders at several localities on the east coast
of northern New Zealand. It occurs throughout New
Zealand and also in Australia.
Berthella ornata (Cheeseman, 1878) Fig. 9
Average animal length 60 mm. This large attractive
species has a base colour of white to cream and is
marked with dark reddish brown blotches. The skin is
smooth and slippery to touch, but there are microscopic
calcareous spicules present. The large gill is long,
extending as far as the tail. There is a small internal shell.
The soft spiral spawn coils have a diameter of up to 15
mm, are peach to cream in colour and attached under
low tidal rocks (pers. obs.).This endemic pleurobranch
is rare intertidally to 6 m, preferring partially sheltered
and open coasts, but not harbours (Willan and Morton
Margaret S. Morley & Bruce W. Hayward
65Intertidal records of ‘sea slugs’
1984). We have found it in low numbers on exposed
and sheltered coasts along the east side of northern New
Zealand with one record from Manukau Harbour on the
north west side. It occurs right around mainland New
Zealand and on the Chatham Islands.
Berthellina citrina (Ruppell and Leuckart, 1828) Fig. 9
Average animal length 50 mm. It is usually orange in
colour though can range from pale yellow to bright red.
The back is covered with small, white gland cells. The
animal has a small internal shell. It has been reported
feeding on sponges (Willan and Morton 1984) and corals
(Sea Slug Forum). We have found occasional specimens
under low tidal stones on moderately sheltered coasts on
the east coast of northern New Zealand and in Manukau
Harbour. It is also known from the Three Kings Islands
and Wellington (Te Papa Museum database) and is
common throughout the Indo-Pacic Ocean.
Pleurobranchaea maculata (Quoy and Gaimard, 1832)
Fig. 10
Average animal length 80 mm. This large species is the
only pleurobranch without an internal shell. It varies from
pale grey to almost black with anastomosing dark wavy
lines. It occurs on all substrates from silty harbours to
open rocky coasts, even to continental slopes in depths of
250 m (Willan and Morton 1984). Although not common,
we have found P. maculata widespread on the east and
west coasts of Northland, and in many localities along the
west side of the Hauraki Gulf and in Waitemata Harbour.
In 2009 it washed up in large numbers on the east
coast bays of Auckland’s North Shore. The population
Figure 11. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Cadlina
willani (Great Barrier, Rangiahua Is), Ceratosoma amoenum (Sentinel Rock, Mangawhai Heads) and Goniobranchus
aureomarginatus (Parengarenga Harbour, Paua). Scale bars 10 mm long.
66
had peaked as a result of feeding on the introduced
Asian mussel, Arcuatula senhousia. Several dogs died
after eating specimens, and subsequent investigations
showed that all parts of the slugs contains a neurotoxin,
tetrodotoxin (McNabb et al. 2010). It is widespread in
temperate waters of the Indian and Pacic Oceans.
Nudibranchia
Cadlinidae
Cadlina willani Miller, 1980 Fig. 11
Average animal length 14 mm. This small nudibranch
is white with a central yellow stripe and border. It has
a rough mantle and the foot extends beyond the body.
It lives among encrusting organisms in open localities
and channels. It is probably the most common subtidal
nudibranch in New Zealand (Willan et al. 2010) and
extremely rare intertidally, with only one such record
from all our studies – Rangiahua Island, o the west
coast of Great Barrier Island (Morley 1991). This
endemic species occurs around the North Island and
south to Banks Peninsula.
Chromodorididae
Ceratosoma amoenum (Cheeseman, 1886) Fig. 11
Average animal length 30 mm. The common name,
clown nudibranch, is well chosen as the ground colour
of the animal is white with vivid orange patches, purple
rhinophores and gill. The details of the design vary
with each individual. The body is soft and slippery.
Cheeseman’s iconotype (MA70169) is from the
Waitemata Harbour. We have found it frequently at many
locations along the east side of northern New Zealand,
with the majority of records from shallow depths of 2–3
m where it appears to be most common. The full depth
range is from low tidal to 40 m on semi-sheltered shores
(Willan and Morton 1984). The range is the northern half
of the North Island to East Cape and Taranaki on the
west coast. It also occurs on Lord Howe and Norfolk
islands and temperate southern Australia.
Goniobranchus aureomarginatus (Cheeseman, 1881) Fig. 11
Average animal length 35 mm. The body is white with
a golden line close to the border of the mantle. It is less
common than Ceratosoma amoenum and rarely found
intertidally where it sometimes crawls exposed on
sponges under low tidal rocks. It lives from low tide
to 24 m on rocky reefs in open situations (Willan et al.
2010). Our records of this endemic species are all from
semi-exposed eastern locations, but it has been recorded
from around the main islands of New Zealand and the
Chatham Islands.
Discodorididae
Alloiodoris lanuginata (Abraham, 1877) Fig. 12
Average animal length 60 mm. This large nudibranch
varies from reddish brown to grey with a pattern of
brown and white spots. The mantle is rough due to minute
tubercles with projecting spicules. The rhinophores are
yellow and the gills greenish with darker markings
(Willan and Morton 1984). The spawn coil is yellowish
tan (pers. obs.). Its habitat is from sheltered to exposed
shores from intertidal down to depths of 10 m where it
feeds on sponges. Single specimens or pairs have been
found at our study sites on both east and west coasts of
northern New Zealand, but not in the Far North. This
endemic species occurs around all the main islands and
the Chatham Islands.
Atagema molesta (Miller, 1989) Fig. 12
Average animal length 20 mm. This nudibranch is fawn
with brown spots and has a soft mantle covered in
spiculose tubercles. Those near the mid-dorsal ridge are
opaque white. It is uncommon and occurs beneath stones
in tide pools on semi-sheltered coasts. It has only been
recorded from Goat Island Bay, the type locality, and the
Whangaparaoa Peninsula.
(Miller 1989, Willan et al. 2010).
Hoplodoris nodulosa (Angas, 1864) Fig. 12
Average animal length 35 mm. This uncommon
nudibranch is yellow, brown or grey with brown blotches,
white pustules, brown gills and rhinophores. There is an
unpigmented area around each pustule. We have found
this at only three study localities – on the sheltered east
and west coasts of Auckland. It was most frequent on the
compound ascidian Alcyonium aurantiacum at Jenkins
Beach, Manukau Harbour. This species also occurs
around the main islands of New Zealand and along the
coast of southern Australia.
Jorunna pantherina (Angas, 1864) Fig. 12
Average animal length 15 mm. A pale peachy bu
nudibranch with marbled ring patches which grade
smaller towards the margins. The colour can vary from
yellow to black according to the sponge eaten (Willan
and Coleman 1984). The gill has overlapping sections
like petals on a rose. The dorsum is thickly beset with
short tubercles. It has been found in just a few shallow,
relatively sheltered locations on the east side of
Auckland and southern Northland. We have found only
one intertidal specimen – at Meola Reef, Waitemata
Harbour on low tidal rocks with a grey sponge. It
is also found in Australia (Sea Slug Forum).
Rostanga muscula (Abraham, 1877) Fig. 12
Average animal length 18 mm. This small nudibranch is
tomato red with scattered dark spots. It is often found at
low tide on its host, a red sponge. It lays a delicate red
spawn coil with a diameter of 12 mm. We have found
this endemic species in low numbers in a wide range of
mid to low tidal semi-sheltered to exposed sites on both
sides of northern New Zealand. It has been recorded as
far south as the Marlborough Sounds (Willan et al. 2010).
Dorididae
Aphelodoris luctuosa (Cheeseman, 1882) Fig. 13
Average animal length 50 mm. A graceful dorid with
a ground colour of cream variably ornamented with
Margaret S. Morley & Bruce W. Hayward
67Intertidal records of ‘sea slugs’
Figure 12. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Atagema
molesta, Alloiodoris lanuginata (Kawau Island, Mansion House Bay), Haplodoris nodulosa (Manukau Harbour, Jenkins
Bay), Jorunna pantherina (Waitemata Harbour, Meola Reef) and Rostanga muscula (Bay of Islands, Tapeka Pt). Scale
bars 10 mm long.
radial brown, wide stripes, mid dorsal blotches and ne
concentric lines around the mantle edge. The rhinophores
and gills are chocolate-brown, rimmed with yellow. The
pale peach spawn coil in a depth of 2 m at Rangiputa
measured 70 mm in diameter (pers. obs.). It crawls
exposed on rocky coasts down to 40 m depth (Willan and
Morton 1984) and is only seen occasionally intertidally.
The unpublished iconotype from the Waitemata Harbour
was painted by Emma Cheeseman (MA70246). We have
records from exposed and sheltered shores on both sides
of northern New Zealand. This endemic species occurs
around the main islands and the Chatham Islands.
Aphelodoris sp. Fig. 13
Average animal length 35 mm. This undescribed dorid
has an encircling pattern of tan blotches on cream but
lacks the dark concentric lines of A. luctuosa. The
gills and rhinophores are white, there is a pinkish tinge
on the underside of the mantle edge (pers. obs.). All
records of this endemic species are from the exposed to
semi-sheltered west coast of northern New Zealand (e.g.
Hayward and Morley 2004).
Doris granulosa (Pease, 1860) Fig. 14
Average animal length 25 mm. A distinguishing feature of
this yellow nudibranch is the posterior facing, horizontal
gill which can be withdrawn under a ap of the mantle.
There are low rounded tubercles over the whole of the
mantle. It is often found feeding on sponges. It lives
from mid to low tide on rocky reefs in semi-protected
localities right around New Zealand. It also occurs in
New Caledonia and temperate Australia.
68
Doris wellingtonensis Abraham, 1877 Fig. 14
Average animal length 100 mm. This is New Zealand’s
largest dorid. The eshy body is covered with rounded
pustules. It is khaki brown to yellowish above, and the
foot and rhinophores are orange (Willan and Morton
1984). We have seen four specimens with three bright
orange spawn masses 30–60 mm diameter in low tidal
pools at Maori Bay, Waitakere coast. This species lives
on exposed to semi-sheltered rocky shores ranging from
intertidal down to 20 m, feeding on yellow encrusting
sponges. It occurs on both sides of northern New Zealand
and around the main islands of New Zealand as well as
eastern Australia.
Dendrodoridae
Dendrodoris citrina (Cheeseman, 1881) Fig. 15
Average animal length 55 mm. This lemon yellow
to orange nudibranch is distinguished from all other
similarly coloured New Zealand species by the presence
of numerous small white dots throughout its length.
The unpublished iconotype (MA70244) is a water
colour painting by Emma Cheeseman. This is the most
commonly encountered and most abundant sea slug in
the intertidal zone of northern New Zealand, where it
occurs on in more shaded locations on mid-low tide
rocky reefs on moderately exposed to harbour shorelines.
It has not been recorded from along the exposed west
coast. It appears to be most common in the inner Hauraki
Margaret S. Morley & Bruce W. Hayward
Figure 13. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Aphelodoris
luctuosa (Waiheke Is, Oneroa) and Aphelodoris sp. (Muriwai, Maori Bay). Scale bars 10 mm long.
69Intertidal records of ‘sea slugs’
Figure 14. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Doris
granulosa (Auckland, Te Arai Pt) and Doris wellingtonensis (North-west Nelson, Tata Bay). Scale bars 10 mm long.
Gulf, especially along the shores of the middle and
outer Waitemata Harbour and the Tamaki Estuary. This
endemic species is recorded from around the two main
islands of New Zealand.
Dendrodoris krusensternii (Gray, 1850) Fig. 16
Average animal length 60 mm. Its vibrant colouring
makes this large dorid conspicuous on low tidal rocks
down to depths of 5 m. The ground colour of the mantle
is light brown with large pale protuberances and folds.
Smooth areas between the folds are peacock blue
(Willan and Morton, 1984). The pale mantle edge has
prominent concentric brown lines. In New Zealand this
species occurs on semi-sheltered coasts on the east side
of northern New Zealand, mainly in the inner Hauraki
Gulf and Waitemata Harbour. Since 2010 specimens
have become more common at more southern localities
than previously found, especially in the Waitemata. It
is widespread throughout the tropical and subtropical
Indo-Pacic Oceans.
Dendrodoris nigra (Stimpson, 1855) Fig. 16
Average animal length 40 mm. This smooth black
dorid has rhinophores tipped with white. The edge of
the mantle is frilled. It lays spirals of orange spawn.
Specimens usually are seen intertidally on sheltered and
semi-open rocky shores on the east coast of northern
New Zealand, mainly within the Hauraki Gulf. It is
widespread throughout the temperate waters of the
Indo-Pacic Ocean.
70
Polyceridae
Plocamopherus imperialis Angas, 1864 Fig. 17
Average animal length 70 mm. This is a large dramatic
nudibranch that was rst seen in New Zealand about
50 years ago (M. Miller pers. comm.). The animal is
scarlet with small dark spots and two longitudinal cream
stripes. The rhinophores are a ambuoyant orange with
bulbous purple tips. Posterior to the gills are two globes,
which ash light if the animal is disturbed. The oral
shield has small compound papilli around the margin.
The large brown brachial plume is attached to the body
on a pedestal (Morley 1996). We found this species in
low numbers at Dysons Beach, Parengarenga Harbour.
One specimen was laying an orange spawn coil on a low
tide frond of brown sea weed, Carpophyllum sp. Another
pair was crawling on sand ats in depths of 3 m. When
disturbed the animal turns sideways and swims strongly
with muscular contractions of the body and tail (pers.
obs.). This species is widespread throughout the tropical
and subtropical Indo-Pacic and is recorded here from
the Three Kings Islands and two localities on the east
coast of Northland.
Polycera hedgpethi Er. Marcus, 1964 Fig. 17
Average length 13 mm. This long, narrow species has
a cream body peppered with black. The head has four
elongate papilli and there is a cluster of two or three papilli
beside the gills. These are translucent white with a broad
gold ring near the centre and a dark zone above. The tip
of the tail is also gold (Willan et al. 2010, p.438). It is
recorded from low intertidal to 10 m depth but mostly
Margaret S. Morley & Bruce W. Hayward
Figure 15. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Dendrodoris
citrina (Waiwera). Scale bar 10 mm long.
71Intertidal records of ‘sea slugs’
Figure 16. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Dendrodoris
krusensternii (Kohimarama, Waitemata Harbour) and Dendrodoris nigra. Mellons Bay). Scale bars 10 mm long.
from the sublittoral fringe. It was probably introduced into
New Zealand in the 1960s living on bryozoa attached to
a ship’s hull (Willan et al. 2010). It is recorded from the
Waitemata Harbour, Cavalli Islands and the Marlborough
Sounds. It is cosmopolitan in warm temperate seas.
Polycera melanosticta Miller, 1996 Fig. 17
Average animal length 14 mm. This is the species
recorded in Morton & Miller (1968) as Polycera sp. The
body is elongated bearing tubercles on the head veil, gill
circlet and variously along the body. It is white, grey or
greyish brown patterned with white, black and orange
spots widely spaced over the body (Miller 1996). It was
formally described by Miller (1996) from a number of
specimens collected from a paint raft at the type locality
at Devonport naval base, Waitemata Harbour in 1962. It
may have been introduced to New Zealand on a ship’s
hull. It also occurs in Victoria, Australia (Burn 2006).
Thecacera pennigera (Montagu, 1815) Fig. 17
Average animal length 12 mm. A small high bodied
nudibranch with a deep pit in front of each rhinophore.
The rhinophores are surrounded by anged sheaths and
two nger-like processes arise behind the gills. Each
individual has a unique pattern of orange and black
spots on the white body. It was recorded commonly in
the upper Waitemata Harbour and Tamaki Estuary in the
1970s, but has not been seen more recently. It was found
on hard substrate in the sublittoral fringe and subtidal
where it feeds on bryozoa. It is presumed to have been
introduced by shipping (Willan 1976) but may not have
survived (Willan et al. 2010).
72
Proctonotidae
Caldukia rubiginosa Miller, 1970 Fig. 17
Average animal length 10 mm. This small reddish-brown
nudibranch has cerata on the body and around the head.
The upper part of each ceras is yellow and the central
section iridescent blue. It is occasionally found among
encrusting organisms under stones in the low intertidal
and sublittoral zones down to 13 m on open coasts
(Willan et al. 2010). It was not found in our surveys but
this endemic species has been recorded from the Three
Kings Islands and sporadically along the east coast of
northern New Zealand as far south as Hahei, Coromandel
Peninsula. The type locality is Goat Island Bay, Leigh.
Janolus novozealandicus (Eliot, 1907) Fig. 17
Average animal length 20 mm. The translucent body
has opaque white marks down the centre and brown and
white speckles over the sides. Inside each ceras can be
seen the orange and blue digestive diverticulum (Willan
et al. 2010). This endemic species is found beneath
stones on exposed to semi-sheltered reefs and channels
at low tide to depths of 15 m along the east coast of
northern New Zealand, and has been recorded around
all three major islands (Miller and Willan 1986). It is
uncommon, but more frequent in South and Stewart
Islands (Willan et al. 2010).
Margaret S. Morley & Bruce W. Hayward
Figure 17. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Plocamopherus
imperialis (Parengarenga, Dysons Beach), Polycera hedgpethi, Polycera melanosticta, Thecacera pennigera, Caldukia
rubiginosa (redrawn after Willan and Morton, 1984, p. 68; Leigh, Goat Island Bay), Janolus novozealandicus and Vayssierea
cinnabarea (Northland, Kawerua). Scale bars 10 mm long unless stated otherwise.
73Intertidal records of ‘sea slugs’
Figure 18. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Fiona
pinnata (redrawn after Willan and Morton, 1984, p. 81), Acanthodoris mollicella (Manukau Harbour, Kaitarakihi Bay),
Goniodoris n.sp. (Bay of Islands, Deep Water Cove), Okenia plana (redrawn after Willan and Morton, 1984, p. 91) and
Trapania rudmani (Parengarenga, Paua). Scale bars 10 mm long unless stated otherwise.
Vayssiereidae
Vayssierea cinnabarea (Ralph, 1944) Fig. 17
Average animal length 4 mm. This small slender dorid
nudibranch is dull red, it has no gill or retractable
rhinophores (Willan and Morton 1984). It lives on the
undersurface of intertidal stones on open rocky coasts in
the presence of Spirorbis tube worms on which it feeds
(Willan and Morton 1984). In this study this endemic
species is recorded from one locality on either side of
northern New Zealand. It is probably more common than
this but overlooked due to its small size. Its type locality
is Wellington.
Fionidae
Fiona pinnata (Eschscholtz, 1831) Fig. 18
Average animal length 17 mm. This pelagic nudibranch
is usually cream or bu in colour with an orange patch
between the rhinophores. It becomes blue when feeding
on by-the-wind sailor, Velella velella and Portuguese
man-of-war, Physalia physalis. The cerata are in oblique
rows with a membranous attachment on the inner margin.
There are pairs of rhinophores and oral tentacles. It
attaches to oating wood or Durvillaea kelp which host
the barnacles Lepas spp. F. pinnata can feed on several
dierent species of Lepas (Willan 1979b). Amazingly for
a pelagic species F. pinnata cannot swim. The species
74
has been found occasionally washed up on open coasts
following prolonged on-shore winds, right around New
Zealand including the Chatham and Subantarctic islands.
It is cosmopolitan in tropical and temperate seas.
Onchidorididae
Acanthodoris mollicella Abraham, 1877 Fig. 18
Average animal length 25 mm. This small nudibranch is
cryptically coloured in mottled black, brown and ochre
yellow. The dorsum is covered in numerous papilli,
and it has transparent feathery gills which tend to point
backwards. The tips of the rhinophores are white. This
endemic species lives in protected channels and harbours.
We have found solitary specimens under low tidal rocks
in Manukau, Waitemata and Whangarei Harbours. It is
also known from around most of New Zealand.
Goniodorididae
Goniodoris n.sp. Fig. 18
Animal length 10 mm. This species has an unusual
exposure of the tail and is only known from a single
specimen found under a rock in depth of 2 m at low tide
in Deep Water Cove, Bay of Islands. The specimen is
preserved in ethanol and its original colour is unknown. A
second species, G. castanea, is also represented by only
one specimen, dredged in Otago Harbour, South Island.
This North Island record may prove to be a subspecies G.
castanea (Michael Miller pers. comm.). Goniodoris spp.
are also found in North America and Japan.
Okenia plana Baba, 1960 Fig. 18
Average animal length 8 mm. This nudibranch has a body
copiously spotted with lilac and reddish-brown. It has
a broad foot, large rhinophores and a mantle edge with
a row of tentacle-like projections. Its habitat is among
brown algae and bryozoans in protected situations and
silty harbours (Willan and Morton 1984). It has been
recorded in the Waitemata Harbour and at Leigh, but has
not been found in our studies. It is also found in Australia,
Hawaii, Japan and Malaysia (Sea Slug Forum).
Trapania rudmani Miller, 1981 Fig. 18
Average animal length 18 mm. The body of this slender
nudibranch gradually tapers to a long tail. There are two
pairs of dorso-lateral nger-like processes. The body is
translucent white with yellow stripes on the outside of
the four processes. It is rare intertidally and often lives
on vertical walls among encrusting organisms in open
localities down to depths of 20 m (Miller 1981). We
record it here from wharf piles at 1 m depth at Paua,
Parengarenga Harbour, Northland. This endemic species
has been recorded subtidally along the east coast of
northern New Zealand (Sea Slug Forum).
Arminidae
Dermatobranchus rubidus (Gould, 1852) Fig. 19
Average animal length 20 mm. The mantle is dark orange
while the longitudinal ridges, rhinophores and oral veil
are highlighted in white. The foot is wider than the
mantle and has tentacles at the front corners (Willan et
al. 2010). The species lives in semi-sheltered localities
on sand from the sublittoral fringe to 65 m. Only two
specimens of this rare nudibranch were previously known
from New Zealand. We have found a third specimen in a
depth of 2 m at Otupoho Bay, Bay of Islands. There is a
record from McDonald Bank, Whangarei Harbour. It is
also found in Australia.
Tritoniidae
Tritonia incerta Bergh, 1904 Fig. 19
Average animal length 100 mm. This large uncommon
nudibranch has three colour forms, pale rose-pink,
apricot orange and red with white gills and foot. The gills
consist of tufts along each side of the notal ridge and the
oral veil has ten or twelve tentacles (Willan et al. 2010).
The depth range this endemic species extends down to
250 m. It has been recorded intertidally from the exposed
Waitakere coast at Maori Bay, Muriwai (Powell 1979,
p. 287) and Whatipu, Manukau Heads. It has not been
found it in our systematic surveys. It also occurs further
south in New Zealand on both the east and west coasts of
the North Island and the west coast of the Soiuth Island as
far south as Westport (Te Papa Museum database).
Dotidae
Doto pita Er. Marcus, 1955 Fig. 19
Average animal length 8 mm. This small slender
nudibranch has up to seven pairs of nger-like cerata
on either side of its body. The body is translucent white
mottled with yellow, brown and lilac spots (Willan et
al. 2010). It is mostly found feeding on hydroids and
can be common at low tide on open rocky reefs and in
semi-sheltered locations. It also extends to depths of
30 m. It is known in New Zealand from the east coast,
north of Auckland and elsewhere from Brazil, Japan and
Australia (Willan et al. 2010).
Flabellinidae
Flabellina albomarginata (Miller, 1971) Fig. 19
Average animal length 18 mm. This species has a
transparent whitish body with a white border on the foot,
and red cerata tipped with white. It lives from intertidal
down to depths of 20 m in open to semi-sheltered
localities. In northern New Zealand this endemic species
has been found intertidally on the Waitakeres’ west coast
between Whatipu and Muriwai, but not on our systematic
surveys. The type locality is Piha. This endemic species
is more common on the Canterbury and Otago coast of
the South Island.
Tularia bractea (Burn, 1962) Fig. 19
Average animal length 15 mm. This nudibranch has a
white to yellowish brown body with pointed corners to the
anterior of the foot, long rhinophores and oral tentacles.
The seven clusters of cerata arise from brackets. The red
spots along the edge of the cerata perfectly match the
Margaret S. Morley & Bruce W. Hayward
75Intertidal records of ‘sea slugs’
Figure 19. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of
Dermatobranchus rubidus (Bay of Islands), Flabellina albomarginata (Waitakere coast, Piha), Doto pita, Tularia bractea
(Waitemata Harbour, Musick Pt), Eubranchus agrius (after Willan and Morton, 1984, p. 79), Trinchesia beta, Trinchesia
refl xa and Tritonia incerta (Waitakere coast, Whatipu). Scale bars 10 mm long unless stated otherwise.
lolly-pink of coralline algae on which we have found it
living. It lives intertidally down to depths of 25 m and is
more common subtidally. We have found one specimen
at Musick Point, Tamaki Estuary on coralline algae in
mid-tidal pools. It occurs around both North and South
Islands and is also known from Victoria, Australia
(Willan et al. 2010).
Eubranchidae
Eubranchus agrius Er. Marcus, 1959 Fig. 19
Average animal length 6 mm. The body of this small
transparent nudibranch has small patches of gold, yellow,
brownish-orange, pale ruby or purplish red. It lives on
hydroids in tidal pools on open coasts. We have not
76
found this species in our studies, but it has been reported
from both the west and east sides of the Auckland region
and occurs around both main islands of New Zealand. It
is also known from Chile.
Tergipedidae
Trinchesia beta (Baba & Abe, 1964) Fig. 19
Average animal length 6 mm. The wide foot extends
beyond the visceral part of the body. The anterior corners
have arc-shaped anges. There are eleven rows of cerata.
The body is translucent, nely spotted with purple. The
cerata are also purple except for a gold or orange band
below the translucent tips (Miller 1977). It has been
recorded in tidal pools, under stones, and on hydroids
in the sublittoral fringe from Whangarei Heads to the
Waitemata Harbour on the east coast and the Waitakeres
on the west (Miller 1977). It was possibly more common
in the 1950s-1960s than it was around northern New
Zealand in the 1990s-2000s, as we did not see it in our
surveys. It is also known from Japan (Miller 1977).
Trinchesia reexa (Miller, 1977)
Average animal length 7 mm. The foot is wide,
extending beyond the visceral part of the body. The
anterior corners are produced as semicircular processes
(Miller 1977). There are nine rows of cerata. The body
is transparent or pale yellow with the viscera clearly
visible. The diverticula are fawn to black. It is usually
found living on hydroids in mid-low tide pools on
Margaret S. Morley & Bruce W. Hayward
Figure 20. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Aeolidiella
drusilla (Northland, Whangamumu), Anteaeolidiella lurana (Parengarenga, Dog Is), Baeolidia australis (Bay of Islands, Tapeka
Pt), Burnaia helicochorda and Phidiana milleri (Oneroa, Waiheke Island). Scale bars 10 mm long unless stated otherwise.
77Intertidal records of ‘sea slugs’
exposed rocky shores. This endemic species has been
recorded from the Waitakere west coast and Manukau
Harbour (type locality); and further south on Banks and
Otago Peninsulas in the South Island (Miller 1977).
Aeolidiidae
Aeolidiella drusilla Bergh, 1900 Fig. 20
Average animal length 30 mm. The body of this
nudibranch is bu with white blotches. The brown and
orange cerata are tipped with orange, and the papillose
grey-brown rhinophores are white-tipped (MSM pers.
obs.). Although considered rare (Powell 1979, p 289 as A.
faustina) thirty to forty specimens were seen at low tide
on ascidians on Paua wharf piles, Parengarenga Harbour
in 1992. We also record a pair found under a rock in 1
m depth at Whangamumu, east Northland. It has also
been found around the main New Zealand islands and the
Chatham Islands, and it occurs in south-eastern Australia.
Anteaeolidiella lurana (Bergh, 1888) Fig. 20
Average animal length 15 mm. Individuals vary from
orange to brown with orange stripes on the head and
a series of white blotches down the back. It lives from
mid-tide to depths of 11 m on rocky reefs in semi-sheltered
localities. We have found it curled up with bryozoa under
low tidal stones at Dog Island, Parengarenga Harbour.
All our records are from the east side of northern New
Zealand as far south as Birkdale and West Tamaki Head,
Waitemata Harbour (Miller 1977). It is widespread in
tropical and subtropical seas and is possibly an introduced
species to New Zealand (Willan et al. 2010).
Baeolidia australis (Rudman, 1982) Fig. 20
Average animal length 20 mm. This colourful nudibranch
has a brown body with numerous white spots. The white
tipped cerata are banded with brown, crimson and gold.
It can be identied by the white hexagonal marking on
the head between the oral tentacles. The cerata are large
and bulbous. It is found on open coasts on seaweeds
where it likes to feed on the anemone Cricophorus
nutrix (Willan & Morton 1984). We have one record
from Tapeka Point, Bay of Islands on the brown seaweed
Carpophyllum. It is also found in Australia.
Burnaia helicochorda (Miller, 1988) Fig. 20
Average animal length 20 mm. The body is translucent
brown with horseshoe-shaped clusters of cerata.
A distinctive feature is the pair of large club-like
rhinophores. Only two specimens have been found in
New Zealand, one at the Three Kings Islands and one
at the type locality Goat Island Bay, Leigh. The habitat
is on exposed rocky shores at low tide to 20 m depth
(Willan et al. 2010). It also occurs in Australia.
Facelinidae
Phidiana milleri Rudman, 1980 Fig. 20
Average animal length 30 mm. An attractive nudibranch
that is frequently noticed because it moves rapidly and
often crawls on exposed surfaces. The slender, translucent
body has orange around the head and tentacles. The
orange cerata have white tips and are darkened centrally
by branches of the digestive gland. This endemic species
is predominantly intertidal, frequently found in intertidal
pools, but extends to depths of 21 m (Willan and Morton
1984). We have scattered records from the east coast of
Northland and it is more common around the Waitemata
Harbour and on the Waitakere west coast. It occurs right
around the North Island and south to Banks Peninsula.
Babakinidae
Babakina caprinsulensis (Miller, 1974) Fig. 21
Average animal length 15 mm. The body is pale mauve
with an iridescent white rectangle on the head. The oral
tentacles are white tipped with yellow. The club shaped-
rhinophores arise from a common pedicle and are
orange-brown also tipped with yellow. There is a narrow
white line on the tail (MSM pers. obs.). The cerata are
white, tipped with yellowish green (Miller 1974, as
Babaina). All records of this endemic species are from
the east coast of northern Northland. The type locality
of this rare species is Goat Island Bay, Leigh. A second
specimen was found under a stone in an intertidal pool
at nearby Mathesons Bay (Morley 1987). We know of
records from Tapeka Point, Bay of Islands in a depth of
17 m where a specimen was spawning, and Whangaroa
Harbour, Northland (Kevin Burch pers. comm.).
Glaucinidae
Glaucus atlanticus Forster, 1777 Fig. 21
Average animal length 20 mm. This pelagic nudibranch
amazes with its glowing colours of blue, silver and
purple. The cerata are clustered on three pairs of lateral
lobes, which assist its buoyancy. It oats upside-down
on the surface of the water and remains aoat by
periodically gulping air into its stomach. It feeds on
Portuguese man-of-war, Physalia physalis, by-the-wind
sailor Velella velella and the hydrozoan Porpita porpita
(Willan and Morton 1984). It has been recorded washed
up on both coasts of northern New Zealand. It is
seasonally frequent in the Kermadec Islands and occurs
world-wide in tropical and warm temperate zones.
Umbraculidae
Umbraculum umbraculum (Lightfoot, 1786) Fig. 21
Average animal length 120 mm. This uncommon
side-gilled sea slug has a massively thiumbraculuckened
body covered in tubercles. The body colour varies from
grey to brown, yellow, reddish and orange (Willan et
al. 2010). The attened, oval shell perches on top. Its
habitat is on open rocky reefs from intertidal down to
40 m. It has been found on the east coast of northern
New Zealand as far south as East Cape (Te Papa
Museum database). It is also found in all tropical and
subtropical seas.
78
Velutinidae
Lamellaria ophione Gray, 1850 Fig. 22
Average animal length 18 mm. Although the soft body
of this species is similar externally to a nudibranch it
is not related (Powell 1979, p.150). The smooth body
varies in colour from yellowish, orange, grey to white,
to blend with its prey of ascidians and sponges. It has
a fragile internal shell which is occasionally found in
shell sand. This endemic species lives under rocks at
low tide on exposed to sheltered coasts and is frequently
seen on both sides of northern New Zealand. It appears
to be most common in the Manukau Harbour, especially
at Jenkins Bay, where it was found on the compound
ascidian Alyconium aurantium. We know of one South
Island record from Haast (Te Papa Museum database).
Onchidiidae
Onchidella nigricans (Quoy & Gaimard, 1835) Fig. 23
Average animal length 20 mm. The leathery sea slug is
a pulmonate, it has no shell and as the common name
suggests, has a tough granulated outer skin, often with
papilli. It varies in colour from black to grey and maybe
mottled with white, black, orange, green or yellowish-
green (Willan et al. 2010). There are white marginal
glands. It secretes a noxious uid to deter predators.
We have found it widespread in our studies on both
coasts of Northland where it is most common on shaded
muddy surfaces from mid to low tide. Occasionally on
semi-exposed beaches it seeks shelter in disused isopod
burrows (pers. obs.). It grazes on diatoms and other
organic debris (Morton & Miller 1968, p 830). It is
Margaret S. Morley & Bruce W. Hayward
Figure 21. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Babakina
caprinsulensis (Leigh, Mathesons Bay), Glaucus atlanticus (Northland, Spirits Bay) and Umbraculum umbraculum (Bay of
Islands, Long Beach). Scale bars 10 mm long.
79Intertidal records of ‘sea slugs’
found throughout New Zealand and also occurs on the
southeast coast of Australia and Tasmania (Willan et
al. 2010).
Fissurellidae
Scutus breviculus (Blainville, 1817) Fig. 24
A mollusc specialist would not class this as a ‘sea-slug’,
but to the layperson it has a slug-like appearance and so
we have included it here for completeness.
Average animal length 120 mm. The velvety black
slug-like animal has a attened, thickened shell which is
almost entirely concealed by the mantle (Powell 1979).
This shell is shield shaped and protects the heart and
other vital organs. The eyes are at the outer bases of long
tentacles. It can be found ocasionally under boulders,
in deep crevices or overhangs at low tide on open to
semi-exposed beaches. It is widespread throughout our
study area on the east and west coasts. It is endemic,
recorded from North, South and Chatham Islands.
DISCUSSION
Several small species of sea slug (less than 10 mm
across) that have been recorded by others from the
intertidal or shallow subtidal of northern New Zealand
(e.g. Okenia plana, Caldukia rubiginosa) were not
found in our surveys as their specic habitats were not
targeted. Had they been present they may have gone
unnoticed on account of their small size.
Figure 22. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Lamellaria
ophione (Whangaparaoa Peninsula, Red Beach). Scale bars 10 mm long.
80
Sporadic abundances
Our surveys conrm the observations of others about the
unpredictability of the presence and size of populations
of sea slugs at any one time or place (e.g., Enderby
and Enderby 2005). The most frequently encountered
intertidal nudibranch around northern New Zealand is
Dendrodoris citrina, often as closely associated pairs or
in numbers fewer than 20 on any one shore (pers. obs.).
Occasionally there have been short periods when
large numbers of one sea slug species has been present
along a short section of coastline. This has been the
case for the green sea hare Bursatella leachii which has
been seen in large numbers at intertidal and shallow
subtidal depths on a number of occasions, often with
wash ups of dying seaweed and cyanobacteria – e.g.,
Orakei Basin, autumn 1996 (Hayward and Hayward
1999); Blackpool and Whakanewha, Waiheke Island,
March 1999 (MSM pers. obs.); Beachlands to the
North Shore, Auckland, March-July 2000 (Morley et
al. 2001); Tindalls Beach, Whangaparaoa, Feb 2015
(this study). The well-camouaged B. leachii are rst
noticed by their bright green spawn, like tangled balls
of knitting wool, which may lead to the discovery of
the recently dead bodies of this sea hare, as they die
after spawning (Morton and Miller 1968). Sea hares of
the genus Aplysia are also known to occur sometimes
in large numbers intertidally – e.g., A. argus and A.
keraudreni, Mathesons Bay and Echinoderm Reef,
Goat Island, Feb-March 1992 (A. Enderby, pers. comm.
2015); A. parvula grazing on brown algae at Leigh
Marine Reserve in 1991, 1997, 1998, and 2004 (A.
Enderby, pers. comm. 2015). Another sea slug to occur
periodically in large numbers and that may result in dog
poisonings when eaten by them, is Pleurobranchaea
Margaret S. Morley & Bruce W. Hayward
Figure 23. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Onchidella
nigricans (Waitemata Harbour, Eastern Beach). Scale bars 10 mm long.
81Intertidal records of ‘sea slugs’
Figure 24. Intertidal and shallow subtidal (down to 2 m) distribution records around northern New Zealand of Scutus
breviculus (Waitemata Harbour, Stanley Bay). Scale bars 10 mm long.
maculata (Khor et al. 2014), which had a major increase
in numbers along Auckland’s North Shore beaches (pers.
obs.) in the summer of 2009-2010 possibly as a result of
an abundance of Asian date mussels just oshore (W.
Blom pers. comm.). One other sea slug to be reported in
large numbers is Tritonia incerta feeding on dead-man’s
ngers, Alcyonium auranticum, at Mathesons Bay and
along the Leigh Marine Reserve coast in Feb 1992-1993
and again in Feb 2004 (A. Enderby, pers. comm. 2015).
Biodiversity
A total of 78 intertidal and shallow marine ‘sea slug’
species are recorded here from northern New Zealand. The
majority are named but three rare taxa (e.g. Aphelodoris sp.,
Elysia n.sp., Goniodoris n.sp.) are probably undescribed
new species. All species, except for Goniodoris n.sp., have
been recorded from New Zealand previously.
Many other unnamed species of opisthobranch
gastropods from New Zealand are recorded in Spencer
et al. (2009). Of the 78 sea slug species, only three
(Aphelodoris sp., Flabellina albomarginata, Trinchesia
reexa) have no intertidal record from the east coast
of northern New Zealand and just 34 (44%) of the
species are recorded from the west coast. The highest
diversities from smaller geographic areas are 47 from
around Leigh; 42 from the Bay of Islands; 38 from Great
Barrier Island; 31 from Waitemata Harbour; 27 from
Parengarenga Harbour; 24 from the Manukau Harbour;
19 from the Waitakere coast and 20 from Waiheke
Island. The greater diversity in some areas reects the
intensity of survey (Leigh, Waitemata and Manukau
Harbours, Waitakere coast) or diversity of habitats
on the warmer east coast (Leigh area, Bay of Islands,
Great Barrier and Parengarenga). Not surprisingly the
82
lowest diversities of sea slugs are around the sediment-
dominated inner reaches of the major harbours and
Firth of Thames where there is reduced habitat diversity
(fewer hard rock substrates, seaweeds and sponges) and
often a smothering with ne sediment.
Biogeography
Thirty-ve species (45%) of the sea slugs recorded here
from northern New Zealand are apparently endemic
to New Zealand. Seven of these endemic species are
restricted in their known occurrence to the north-east
coast of northern New Zealand (Aupourian Province;
Powell 1955). The next largest biogeographic groups
are those whose distribution is Australasian (24%),
cosmopolitan (18%) and IndoPacic (8%). A total of
39 species (50%) are restricted in their New Zealand
occurrence to northern New Zealand (this study
area), possibly including the Kermadec and Three
Kings islands. Twenty-seven species (35%) of the sea
slugs recored here are restricted in their New Zealand
occurrence to the warmer Aupourian or Aupourian and
Kermadecian provinces. A similar number (25 species)
of sea slugs recorded here occur all around the main
islands of New Zealand (Aupourian, Cookian and
Forsterian provinces).
Many sea slugs appear to be well adapted to hitching
lifts on boats and in the last few centuries many have
extended their natural range by successfully establishing
new populations in far o waters. In a number of instances
it is dicult to be sure which sea slugs are native to
New Zealand and which may be recent invasives. An
example of denite human-assisted introduced sea slugs
to northern New Zealand is Dolabrifera brazieri and
possible human-assisted introduced are Anteaeolidiella
lurana and Polycera melanosticta. At least one of
our endemic species (Philine auriformis) has been
introduced overseas (California, USA; Krug et al. 2012).
ACKNOWLEDGEMENTS
We thank Richard Willan and Michael Miller for their
long-standing encouragement, for providing papers and
assisting with tricky identications, and Bruce Marshall
for his identication of Philine species and other
assistance. Thanks to Anna Berthelsen, Kevin Burch,
Andrew Crowe, Tony Enderby, Jack Grant-Mackie, Lyn
Hellyar, Virginia Toy and Richard Willan for providing
additional records and information. We thank a number
of collectors – Charlie Bedford, Brady Doak, Lance
Everitt, Charles Fleming, R.B. Taylor, Miss Wakelin,
Martyn Walker, and Keith Wise, whose specimens are
held in the collection of Auckland Museum and have
provided additional records for this paper. In addition
we thank Wilma Blom, Roger Grace, Hugh Grenfell,
Severine Hannam, Glenys Hayward, Neville Hudson,
Chris Horne, Linda Morley, Gordon Nicholson, Peter
Poortman, Jenny Raven, Jenny Riley, Nancy Smith,
Glenys Stace, Brett Stephenson and Richard Willan,
who have accompanied us in the eld and contributed to
the records. Thanks to Wilma Blom for assistance while
accessing the Auckland Museum data base. Wilma and
John Early reviewed the manuscript and suggested many
improvements. Tony Enderby is thanked for sharing his
intertidal observations on large abundances of various
sea slug species. The manuscript has greatly benetted
from the critical reading by Richard Willan.
REFERENCES
Burn, R., 2006. A checklist and bibliography of the
Opisthobranchia (Mollusca: Gastropoda) of Victoria and
the Bass Strait area, south-eastern Australia. Museum
Victoria Science Reports 10: 1–42.
Enderby, A. and J. Enderby, 2005. New Zealand’s endemic
nudibranchs. Index Vita Malacologica 3: 1–10.
Hayward, C.M. and B.W. Hayward, 1999. Human impact on
Orakei Basin, Auckland. Tane 37, 137–152.
Hayward, B.W. and M.S. Morley, 2004. Intertidal life around
the coast of the Waitakere Ranges, Auckland. Auckland
Regional Council Technical Publication 298, 102 pp.
Hayward, B.W. and M.S. Morley, 2008. Intertidal life of the
Tamaki Estuary and its entrance, Auckland. Auckland
Regional Council Technical Publication 373, 71 pp.
Hayward, B.W., Morley, M.S., Riley, J.L., Smith, N. and G.
Stace, 1995. Additions to the Mollusca from Kawerua,
North Auckland. Tane 35: 183–193.
Hayward, B.W., Morley, M.S., Stephenson, A.B., Blom,
W.M., Grenfell, H.R., Prasad, R., Rogan, D., Thompson,
F., Cheetham, J. and M.Webb, 1999. Intertidal and
subtidal biota and habitats of the central Waitemata
Harbour. Auckland Regional Council Technical
Publication 127, 40 pp.
Hayward, B.W., Stephenson, A.B., Morley, M.S., Blom,
W.M., Grenfell, H.R., Brook, F.J., Riley, J.J., Thompson,
F. and J.J. Hayward, 2001. Marine biota of Parengarenga
Harbour, Northland, New Zealand. Records of Auckland
Museum 37: 45–80.
Hayward, B.W., Morley, M.S., Grenfell, H.R., Carter, R.,
Hayward, G.C., Blom, W.M., and D. Rogan, 2004.
Intertidal biota and washup at Ahipara and Herekino,
Northland west coast. Poirieria 30: 13–25.
Hayward, B.W., Morley, M.S., and H.R. Grenfell, 2012.
Marine biota of the Whananaki coast, east Northland,
New Zealand. Records of Auckland Museum 48: 77–96.
Hayward, B.W., Morley, M.S., Stephenson, A.B., Grenfell,
H.R. and G.C. Hayward, 2014. Intertidal and shallow
subtidal biota of Whangapoua, north-east Coromandel
Peninsula. Poirieria 38: 2–12.
Khor, S., Wood, S.A., Salvitti, L., Taylor, D.I., Adamson,
J., Mcnabb, P. and S.C. Cary, 2014. Investigating the
diet as a source of tetradotoxin in Pleurobranchaea
maculata. Marine Drugs 12: 1–16.
Krug, P.J., Morley, M.S., Asif, J., Hellyar, L.L. and Blom,
W.M., 2008. Molecular conrmation of species status
for the rare Cephalaspidean Melanochlamys lorrainae
(Rudman, 1968), and comparision with its sister species
M. cylindrica Cheeseman, 1981. Journal of Molluscan
Studies 74: 267–276.
Krug, P.J., Asif, J.H., Baeza, I, Morley, M.S., Blom, W.M. and
Gosliner, T.M., 2012. Molecular identication of two
species of the carnivorous sea slug Philine, invaders of
the US west coast. Biological Invasions 14: 2447–2459.
Margaret S. Morley & Bruce W. Hayward
83Intertidal records of ‘sea slugs’
McNabb, P., Selwood, A.I., Munday, R., Wood, S.A., Taylor,
D.I., Mackenzie, L.A., van Ginkel, R., Rhodes, L.L.,
Cornelisen, C., Heasman, K., Holland, P.T. and C. King,
2010. Detection of tetrodotoxin from the grey side-gilled
sea slug – Pleurobranchaea maculata, and associated
dog neurotoxicosis on beaches adjacent to the Hauraki
Gulf, Auckland, New Zealand. Toxicon 56: 466–473.
Miller, M.C., 1974. Aeolid nudibranchs (Gastropoda:
Opisthobranchia) of the family Glaucidae from New
Zealand waters. Zoological Journal of the Linnean
Society 54: 31–61.
Miller, M.C., 1977. Aeolid nudibranchs (Gastropoda:
Opisthobranchia) of the family Tergipedidae from New
Zealand waters. Zoological Journal of the Linnean
Society 60: 197–222.
Miller, M.C., 1981. Trapania rudmani, a new dorid
nudibranch (Gastropoda: Opisthobranchia) from New
Zealand. New Zealand Journal of Zoology 8: 5–9.
Miller, M.C., 1989. Trippa molesta, a new dorid nudibranch
(Gastropoda: Opisthobranchia) from New Zealand. New
Zealand Journal of Zoology 16: 243–250.
Miller, M.C., 1996. A new species of the dorid nudibranch
genus Polycera Cuvier, 1816 (Gastropoda:
Nudibranchia) from New Zealand. Journal of Molluscan
Studies 62: 443–450.
Miller, M.C. and Rudman, W.B., 1968. Two new genera
and species of the superfamily Runcinoidea (Mollusca
Gastropoda: Opisthobranchia) from New Zealand.
Transactions of the Royal Society of New Zealand
10(19): 183–189.
Miller, M.C. and R.C. Willan, 1986. A review of the New
Zealand arminacean nudibranchs (Opisthobranchia:
Arminacea). New Zealand Journal of Zoology 13:
377–408.
Morley, M.S., 1987. Unconrmed sighting of Babakina
caprinsulensis. Poirieria 15(3): 2–3.
Morley, M.S., 1991. An intertidal record of Cadlina willani.
Poirieria 16(3): 35.
Morley, M.S., 1996. Plocamophorus imperialis (Angas,
1864) (Nudibranchia: Polyceridae: Triophinae).
Poirieria 19: 26–27.
Morley, M.S., 2004. A population of the sea slug Ercolania
felina. Poirieria 30: 9–13.
Morley, M.S. and B.W. Hayward, 1999. Inner shelf Mollusca
of the Bay of Islands, New Zealand, and their depth
distribution. Records of the Auckland Museum 36: 119–138.
Morley, M.S. and B.W. Hayward, 2009. Marine Mollusca of
Great Barrier Island, New Zealand. Records of Auckland
Museum 46: 15–51.
Morley, M.S., Hayward, B.W. and A. White, 2001. Changes
to the intertidal biota 1950’s-2000 at Howick Beach.
Poirieria 27: 4–19.
Morton, J.E. and M.C. Miller, 1968. The New Zealand Sea
Shore. Collins, London. 638 pp.
Powell, A.W.B., 1941. Biological primary types in the
Auckland Museum. Records of the Auckland Institute
and Museum 2: 239–259.
Powell, A.W.B., 1955. New Zealand biotic provinces. Auckland
Museum Conchology Section Bulletin 11: 197–207.
Powell, A.W.B., 1979. New Zealand Mollusca. Collins,
Auckland, 500 pp.
Raven, J., 2000. Observations on live Hydatina. Cookia 11:
16–17.
Rudman, W.B., 1998. (December 18) Bullina lineata (Gray,
1825). [In] Sea Slug Forum. Australian Museum,
Sydney. Available from http://www.seaslugforum.net/
factsheet/bulllinea. Retrieved 9 Sept 2015.
Rudman, W.B., 2004 (Feb 10) How many species of
Hydatina are there? [Message in] Sea Slug Forum.
Australian Museum, Sydney. Available from http://www.
seaslugforum.net/nd/12170. Retrieved 9 Sept 2015.
Spencer, H.G., Marshall, B.A., Maxwell, P.A., Grant-
Mackie, J.A., Stilwell, J.D., Willan, R.C., Campbell,
H.J., Crampton, J.S., Henderson, R.A., Bradshaw, M.A.,
Waterhouse, J.B. and J. Pjeta, 2009. Phylum Mollusca:
chitons, clams, tusk shells, snails, squids, and kin. Pp
161–254 in: Gordon, D.P. (ed.) New Zealand Inventory
of Biodiversity, Vol. 1. Kingdom Animalia. Canterbury
University Press, 566 pp.
Spencer, H.G., Willan, R.C., Marshall, B.A. and T.J. Murray,
2014. Checklist of the recent Mollusca recorded from
the New Zealand Exclusive Economic Zone. http://
www.molluscs.otago.ac.nz/
Willan, R.C., 1976. The opisthobranch Thecacera pennigera
(Montagu) in New Zealand, with a discussion on the
genus. Veliger 18: 347–352.
Willan, R.C., 1979a. The ecology of two New Zealand
opisthobranch molluscs. Unpublished PhD Thesis.
University of Auckland, Auckland, New Zealand. 278 pp.
Willan, R.C., 1979b. New Zealand locality records for the
Aeolid nudibranch Fiona pinnata (Eschscholtz) Tane
25: 141–147.
Willan, R.C., 1981. Rediscovery of Runcinella zelandica
Odhner, 1924 (Opisthobranchia: Runcinacea). National
Museum of New Zealand Records 2(2): 5–8.
Willan, R.C. and N. Coleman, 1984. Nudibranchs of
Australia. Australasian Marine Photographic Index,
Sydney, 56 pp.
Willan, R.C. and J.E. Morton, 1984. Marine Molluscs, Part
2 Opisthobranchia. Leigh Marine Laboratory, 106 pp.
Willan, R.C., Cook, S. de C., Spencer, H.G., Creese, R.G.,
O’Shea, S. and G.D. Jackson, 2010. Phylum Mollusca.
Pp. 296–566 in: De Cook, S. (ed.) New Zealand Coastal
Marine Invertebrates, Canterbury University Press, 640 pp.
Margaret S. Morley, Auckland War Memorial Museum.
Bruce W. Hayward, Geomarine Research, 19 Debron Ave, Remuera, Auckland. b.hayward@geomarine.org.nz
APPENDIX 1: (starts over the page) List of intertidal sites surveyed in northern New Zealand and their
sea slug records. * = Date of observations by MSM other than surveys; ** = observers other than the
authors. Numbers prefi ed with MA are specimens lodged in the Marine Department, Auckland War
Memorial Museum. TP ** = records from the on-line Te Papa Museum database.
84 Margaret S. Morley & Bruce W. Hayward
West Coast (north to south)
Cape Maria van Diemen, Jan 1969**. L. ophione
Twilight Beach, Dec 1983*. R. oruaensis
90 Mile Beach, Jan 1965**. P. auriformis (MA32954)
Ahipara, Nov 1994*, May 1999*, TP**. A. luctuosa, A. argus (MA140688), C. thetidis, D. granulosa (MA91075), L. ophione,
M. cylindrica (MA29498), O. nigricans, P. anis, P. maculata, R. oruaensis (MA140618), S. breviculus
Herekino, May 1999*. B. quoyi, H. zelandiae, L. ophione, P. maculata (MA140737), O. nigricans (MA140533), R. oruaensis
Hokianga, TP**. R. oruaensis
Kawerua, Jun 1993. A. lanuginata (MA79338), Aphelodoris sp. (MA131144), B. quoyi, H. zelandiae, O. nigricans,
P. maculata, S. breviculus, V. cinnabarea
Muriwai-Collins Bay, Aug 1999, Sep 2000*, Jan 2002, Dec 2007*. A. lanuginata (MA118232), Aphelodoris sp., D. granulosa
(MA103379), D. wellingtonensis (MA11824), E. felina, F. albomarginata, G. atlanticus, O. nigricans, P. milleri,
P. angasi, P. auriformis, R. muscula (MA118233), S. breviculus, T. incerta
Powell Bay-Tirikohua Pt, Mar 1998, Aug 2003, May 2008. Aphelodoris sp., D. wellingtonensis, E. felina (MA115243,
MA119044), O. nigricans, S. breviculus
Te Waharoa Bay, Jan 2001. O. nigricans, S. breviculus
Te Henga-O’Neills Bay, Jun 1999, Jun 2001. O. nigricans (MA140833), S. breviculus
Anawhata-Fishermens Rock Pt, Sep 1960**, Sep 1999. E. agrius, O. nigricans, S. breviculus, T. beta, T. reexa
Whites Beach, Mar 2000. E. felina, O. nigricans, S. breviculus
North Piha, Nov 1962**, May 2001. F. albomarginata, O. nigricans, T. reexa
South Piha-Lion Rock, Mar 1998, May 2001, TP**. D. wellingtonensis, E. felina, E. agrius, F. albomarginata, F. pinnata,
G. atlanticus, O. nigricans, P. milleri, T. reexa
Mercer Bay, Feb 1998. O. nigricans
Karekare Beach, Oct 1999. O. nigricans
Whatipu-Paratutae, Jan 2000, Sep 2001, Mar 2006, Sep 2006, Dec 2008. Aphelodoris sp. (MA81258, MA119060)),
E. felina (MA108533), F. albomarginata, H. zelandiae, O. nigricans, P. milleri (MA108532), R. muscula (MA10851),
S. breviculus (MA141950), T. incerta
Boulder Bay, Mar 2002, D. granulosa, O. nigricans, R. muscula, S. breviculus
Destruction Gully-Kaiteke Pt, May 1998, Jul 2000, Apr 2001. A. lanuginata, Aphelodoris sp. (MA103518), D.
wellingtonensis, E. felina (MA117874), O. nigricans, R. muscula, S. breviculus
Huia Bay, Mar 1999*, May 2001, Dec 2001*, TP**. A. lanuginata (MA139473), D. granulosa (MA103381), H. zelandiae,
O. nigricans, P. angasi, R. muscula, S. breviculus
Huia Pt-Kaitarakihi, Feb 1999, Sep 2001. A. mollicella (MA140815), B. quoyi, H. zelandiae, L. ophione (MA108534),
O. nigricans, S. breviculus
Puponga Pt and Te Tau Bank, Nov 1994, May 2000, Aug 2001, TP**. A. luctuosa, D. citrina, L. ophione, M. lorrainae,
O. nigricans, P. angasi (MA90365), R. muscula, S. breviculus
Cornwallis-Mill Bay, Feb 1996, May 1997**, Jul 2000, May 2008. B. ornata, B. quoyii (MA28094), D. citrina (MA102568,
MA136851, MA142813), H. zelandiae, O. nigricans, P. angasi (MA142809) R. muscula (MA102575), S. breviculus
Mill Bay-Lawry Pt, Jul 2000. Aphelodoris sp., D. citrina, M. cylindrica, H. zelandiae, O. nigricans, R. muscula
Big Muddy Creek, Dec 2001. B. quoyi, H. zelandiae, O. nigricans
Laingholm, Kauri Pt, Botel Bay, Sep 2006, Nov 2008, Aug 2013. E. maoria (MA117913), H. zelandiae, O. nigricans
Jenkins Bay, Nov 2008. B. quoyi, D. citrina, H. nodulosa (MA118680), L. ophione, O. nigricans, S. breviculus
French Bay, June 2013. H. nodulosa, H. zelandiae, O. nigricans, S. breviculus
Oatoru Bay, Nov 2014. L. ophione, O. nigricans
Waikowhai, TP**. H. zelandiae, O. nigricans
White Blu, Nov 2010. O. nigricans
Onehunga, Nov 2010, TP**. L. ophione, O. nigricans
Mangere Bridge, Mar 2013. O. nigricans
Ambury Park, Mar 2013, Nov 1994. D. citrina (MA90367), H. zelandiae, O. nigricans
Puketutu Is, Mar 2010. H. zelandiae, O. nigricans
Weymouth, Jun 2014. H. zelandiae, L. ophione, O. nigricans
Clarks Beach, Jan 2014. H. zelandiae, L. ophione, O. nigricans
Wattle Bay, 1995**, Oct 2005*. H. zelandiae, M. lorrainae (holotype MA 71314, paratypes MA73138; MA116833),
O. nigricans, P. angasi
Orua Bay, 1956, Jan 1994**, TP**. F pinnata (MA118314), R. oruaensis (paratype, MA71977), S. breviculus
Big Bay, Feb 2015. O. nigricans, S. breviculus
Hamiltons Gap, Awhitu Peninsula, Mar 2004. O. nigricans
Kariotahi, TP**. P. anis
85Intertidal records of ‘sea slugs’
East Coast (north to south)
Te Werahi, Dec 1983*. U. umbraculum
Spirits Bay, Dec 1950**, Mar 1996*, TP**. A. argus, A. parvula, B. vernicosa, G. atlanticus (MA130217), L. ophione,
R. oruaensis (MA31522)
North Cape, TP**, P. anis
Waikuku Beach, **. U. umbraculum
Parengarenga Harbour, Mar 1986*, Mar, Oct 1992*, Mar 1995**, Mar 1996, Sep 2005. A. drusilla, A. lurana (MA130006),
A. parvula, B. vernicosa, B. lineata (MA16926), B. leachii (MA104386), B. ornata, B. quoyi, C. amoenum, C. thetidus
(MA104435), D. citrina (MA77192), G. aureomarginatus, H. physis, H. zelandiae, H. zonata, L. ophione, M. cylindrica,
M. lorrainae (MA116817), O. nigricans, P. milleri (MA77191), P. angasi (MA73137), P. imperialis (MA104471, MA104385),
P. taronga, R. muscula (MA77193), R. oruaensis (MA130147), S. breviculus, (MA104449), T. rudmani (MA77194)
Rangaunu Bay, Nov 1999*, Feb 2001**, TP**. C. thetidis, G. atlanticus (MA105456), H. physis (MA30942), H. zonata,
L. ophione, O. nigricans, P. auriformis, R. oruaensis
Rangiputa, Jul 1994**, Dec 1996**, Oct 2003. A. luctuosa, H. physis (MA139472), H. zonata, M. cylindrica, U. umbraculum
Cape Karikari, Feb 1978**, Apr 1987**, TP**. A. parvula, B. lineata, B. quoyi, Elysia n.sp., G. atlanticus, S. breviculus
Doubtless Bay, 1994**, TP**. B. ornata, H. physis (MA139472), S. breviculus
Taemaro Bay, **. C. amoenum
Taupo Bay, Apr 1988*. D. brazieri, U. umbraculum
Stephensons Island, TP**. P. angasi
Whangaroa, Aug 1961**, TP**. B. caprinsulensis, C, thetidis, H. physis (MA79668), P. anis, P. auriformis, R. oruaensis
Tauranga Bay, TP**. L. ophione, S. breviculus
Mahinepua, **. B. lineata
Cavalli Islands, Nov 1974**, Jan 1979*, Feb 2004**, TP**. B. lineata, C. thetidis, Elysia n.sp., F. pinnata, P. anis,
T. incerta
Matauri Bay, May 1995. G. atlanticus (MA133953)
Bay of Islands, 1971**, Oct 1975**, 1992, Aug 1993, Oct 1994, Nov 1995, Aug 1996, Nov 1997, Dec 1998, TP**. A. argus
(MA88892), A. lanuginata (MA77697), A. luctuosa, A. lurana (MA130006), A. juliana, B. vernicosa, B. australis
(MA77695), B. caprinsulensis, B. citrina, B. leachii, B. lineata, B. medietas (MA140814), B. ornata (MA94796), B. quoyii
(MA140242), C. amoenum, C. thetidis, D. nigra (MA90186), D. brazieri, D. krusensternii, D. granulosa (MA89607),
D. rubidus (MA90186), D. wellingtonensis, G. aureomarginatus, Goniodoris n.sp., H. physis, H. zelandiae (MA133273),
J. novozealandicus; L. ophione (MA133193), O. nigricans, P. anis, P. angasi, P. auriformis, P. milleri, P. powelli,
P. taronga, P. maculata, R. muscula (MA888831), R. katipoides, R. oruaensis, R. zelandica, S. breviculus, T. incerta
Whangamumu Harbour, Dec 1991*. A. drusilla, A. juliana, D. citrina
Taupiri, Apr 1979. U. umbraculum (MA32634)
Bland Bay, Jul 2003*. A. argus, B. lineata, M. cylindrica, P. anis, R. oruaensis
Whangaruru, TP**. M. cylindrica, P. auriformis, S. breviculus
Whananaki, Dec 2000. A. keraudreni (MA104949), B. citrina (MA105891), C. amoenum, C. thetidus (MA165995),
D. brazieri, D. wellingtonensis, G. aureomarginatus (MA105888), H. zelandiae, L. ophione, O. nigricans, P. powelli,
P. maculata (MA105883), S. breviculus
Matapouri Bay, Feb 2003*, TP**. B. lineata, A. parvula (MA113509), A. lanuginata (MA113511)
Tutukaka, Feb 2003**, TP**. A. luctuosa (MA113513), A. parvula (MA113509), B. lineata, H. physis, S. breviculus,
U. umbraculum
Poor Knights Islands, Dec 1977**, 1980*, Mar 1984**, TP**. B. citrina, C. rubiginosa, C. thetidis, C. willani, D. citrina,
Elysia n.sp., J. novozealandicus, P. dendritica, R. katipoides, R. oruaensis, R. zelandica, T. rudmani
Whangaumu Bay, Dec 1991*, May 2013. Aeolidiella sp., B. quoyi, C. thetidis, D. citrina, R. katipoides
Pataua, Sep 2013 B. citrina, B. quoyi, D. citrina, L. ophione, R. katipoides, S. breviculus,
Taiharuru, TP**. D. brazieri, M. lorrainae
Ocean Beach, TP**. G. atlanticus, T. beta
Whangarei Harbour, 1931**, May 1962**, May 1967**, Apr 2012, TP**. A. lanuginata, A. mollicella (MA114018),
B. citrina, B. leachii, B. lineata, B. quoyi, C. amoenum, D. citrina, D. rubidus, H. zelandiae, J. novozealandicus,
J. pantherina, L. ophione, O. nigricans, P. imperialis (MA79667), P. maculata, R. katipoides, R. muscula (MA114017),
R. oruaensis, S. breviculus, , U. umbraculum
Whangarei Heads, TP**. P. anis
Hen Island, TP**. M. cylindrica, U. umbraculum
Chicken Islands, Jan 1982*. C. thetidis, P. anis
Mokohinau Islands, Dec 1977**, TP**. Elysia n.sp., U. umbraculum
Waipu Cove, May 2013. O. nigricans, S. breviculus
Mangawhai Heads, Mar 2014. C. thetidis, A. juliana (MA120614), O. nigricans, P. maculata, S. breviculus,
Te Arai Pt, Sep 2012, TP**. A. keraudreni, A. luctuosa (MA120094), C. thetidis, D. wellingtonensis, L. ophione, P. anis,
S. breviculus
86 Margaret S. Morley & Bruce W. Hayward
Little Barrier Island, 1992**, TP**. C. thetidis, L. ophione, T. incerta
South Pakiri Beach, Mar 2011. O. nigricans, S. breviculus
Goat Island Bay, Leigh, Oct 1961**, Sep 1972, Sep 1991**, May 1999, TP**. A. argus, A. keraudreni, A. luctuosa,
A. lanuginata, A. molesta, A. parvula, B. caprinsulensis, B. citrina, B. helicochorda, B. leachii, B. ornata, C. amoenum,
C. rubiginosa, C. thetidis, C. willani, D. brazieri, D. granulosa, D. nigra, D. wellingtonensis, E. agrius (MA83724),
E. felina, E. maoria, G. aureomarginatus, J. novozealandicus, J. pantherina, M. cylindrica, O. plana, P. anis,
P. maculata, R. katipoides, R. muscula, S. longicauda, S. smaragdinus, T. incerta, U. umbraculum, V. cinnabarea
Mathesons Bay, 1980s*, Feb 1992**, May 1999**, Jul 2000. A. argus, A. juliana, A. keraudreni, A. molesta, B. vernicosa,
B. caprinsulensis, B. leachii, B. medietas (MA118906), B. ornata, B. quoyi, D. brazieri, D. citrina, D. wellingtonensis,
G. aureomarginatus, L. ophione, M. cylindrica, O. nigricans, P. maculata, S. breviculus, T. incerta, U. umbraculum
Ti Pt-Whangateau Harbour, May 1999**, Mar 2010**, May 2012, Sep 2012, Jan 2013. B. leachii, F. pinnata (MA120397),
H. zelandiae, M. lorrainae, O. nigricans, P. anis, P. maculata (MA30818)
Sth end Omaha Beach, Nov 2010. B. quoyi, C. thetidis, L. ophione, P. maculata, S. breviculus,
Takatu Peninsula, Nov 1977**. J. novozealandicus
Jones Bay to Scow Pt, Tawharanui, Aug 2011. A. lanuginata (MA119611), B. ornata (MA119612), B. quoyi, C. thetidis,
L. ophione, O. nigricans, P. maculata, S. breviculus
Christian Bay, Feb 2014. A. argus, B. citrina, B. quoyi, C. amoenum, D. nigra, H. zelandiae, O. nigricans, P. maculata
West Baddeleys Beach, Oct 2012. B. ornata, B. quoyi, C. amoenum, D. nigra, M. cylindrica, O. nigricans
Kawau Island, 1994*, Sep 2011, TP**. A. argus, A. lanuginata, A. parvula, B. leachii (MA79648), B. quoyi, C. amoenum,
H. nodulosa, L. ophione, O. nigricans, P. anis, P. angasi, P. maculata, S. breviculus
Snells-Algies Bays, Sep 2014. B. vernicosa, O. nigricans, R. muscula
Martins Bay, Sep 2014. B. quoyi, M. cylindrica (MA120699), O. nigricans, P. anis
Scott Pt, Mahurangi Harbour, Jun 2001*, Jun 2011. B. quoyii (MA107140), C. amoenum, D. krusensternii, H. zelandiae,
M. cylindrica, O. nigricans
Opahi, Mahurangi Harbour, Aug 2013. A. argus (MA140688), B. leachii, B. quoyi, D. citrina, L. ophione, O. nigricans,
R. muscula, S. breviculus
Cudlip Pt, Mahurangi Heads, Aug 2005, Jan 2010. B. ornata (MA116813), B. leachii, B. quoyi, D. citrina, D. nigra,
D. granulosa (MA116814)
Te Muri Beach, Jan 2012. A. argus, B. quoyi, H. zelandiae, O. nigricans, P. anis
Wenderholm-Waiwera, Jun 1963**, Sep 1974**, Jun 2001, Aug 2006, Jun 2012. B. ornata (MA107049) B. leachii,
B. quoyi, D. citrina (MA119986), D. krusensternii, D. nigra (MA107050), D. pita, E. felina (MA117912), H. zelandiae,
L. ophione, O. nigricans, R. katipoides (MA117915), S. breviculus, T. bractea
Nth Orewa Beach, Jun 2000. B. quoyi, L. ophione, M. cylindrica, O. nigricans, P. auriformis, S. breviculus,
Red Beach, Jun 2008, Jul 2014. A. lurana (MA118987), D. krusensternii, E. maoria, L. ophione, O. nigricans, P. anis,
R. muscula (MA119020), S. breviculus
Manly-Stanmore Bay Pt, Jun 2005. A. lanuginata, B. quoyi, C. amoenum, D. nigra, D. granulosa
Tindalls Beach, 2002*, Feb 2015. B. leachii (MA110910)
Fishermans Rock-Coalmine Bay, Jul 2006, Jun 2007. B. ornata, B. quoyi, D. citrina, L. ophione, O. nigricans
Army Bay, Whangaparaoa Peninsula, May 1999. A. argus, A. juliana, B. leachii, B. quoyi, D. citrina, D. nigra, M. cylindrica
Tiritiri Island, Apr 1931**, Mar 2011, TP**. A. argus, A. parvula, C. thetidus, D. citrina, E. maoria (MA78488),
G. aureomarginatus, O. nigricans, P. maculata, T. incerta, (MA78598)
East Shakespeare Bay, Feb 2010. B. quoyi, D. krusensternii, D. nigra, O. nigricans, S. breviculus
West Okoromai Bay, Dec 2011. B. quoyi, C. amoenum, D. citrina, D. granulosa (MA119768), H. zelandiae, O. nigricans,
S. breviculus
Matakatia Bay and islet, Aug 2010. B. quoyi, C. amoenum, D. citrina, D. nigra, H. zelandiae, M. cylindrica, O. nigricans,
P. maculata
Little Manly, May 2006. B. quoyi, D. citrina, D. nigra, O. nigricans
Arkles Bay, Oct 2010. B. ornata, B. quoyi, C. amoenum, D. citrina, D. nigra, M. cylindrica, O. nigricans
Long Bay, TP**. L. ophione
Nth end Torbay, Oct 2003, Apr 2001, TP**. B. ornata, B. leachii, D. citrina, D. krusensternii, D. nigra, D. granulosa,
L. ophione, O. nigricans
Murrays Bay-Tarata Pt, Jun 2007. B. quoyi, D. citrina, D. krusensternii, M. cylindrica (MA117883), O. nigricans, R. muscula
Mairangi Bay, *. G. atlanticus
Castor Bay, Feb 1998. B. quoyi, M. cylindrica, S. breviculus
Takapuna Reef, 1931**, Oct 1937**, Oct 1960**, Apr 1997, 1999, Jul 2010, TP**. A. mollicella (MA118714), B. ornata,
D. citrina, D. krusensternii, D. nigra (MA135666), E. maoria (MA70259), M. cylindrica (MA70455), O. nigricans,
P. milleri, S. breviculus, T. beta
Nth Narrow Neck Beach, Dec 1958**, Feb 2009. B. quoyi, D. citrina, D. nigra, L. ophione, O. nigricans, P. maculata,
S. breviculus, T. beta
Takapuna Head, Aug 2006, Feb 2009. A. mollicella, B. quoyi, O. nigricans, S. breviculus
87Intertidal records of ‘sea slugs’
Cheltenham Beach-North Head, 1938**, May 2011, TP**. B. quoyi, D. citrina, D. nigra, H. zelandiae, O. nigricans,
P. angasi, P. auriformis (MA31507), S. breviculus
Torpedo Bay, 2001*, Sep 2005. B. ornata, D. citrina, O. nigricans, P. maculata, S. breviculus
Devonport Naval Base, Jul 1961**, May 1969**. J. novozealandicus, P. melanosticta, T. beta, T. reexa
Stanley Pt, Jun 1998, Nov 2004. B. ornata (MA114106, MA136696), B. quoyi, D. granulosa, O. nigricans, P. maculata,
S. breviculus
Northcote-Birkenhead, Jul 1996, Mar 2010, Jun 2010. A. luctuosa, B. quoyi, D. citrina, H. zelandiae, L. ophione,
M. cylindrica, O. nigricans, S. breviculus
Chelsea-Kauri Pt, Aug 2006. B. quoyi, D. citrina (MA117879), D. nigra, D. granulosa, E. maoria (MA117880), O. nigricans,
P. maculata, S. breviculus
Rangitoto Lighthouse, Mar 2002. O. nigricans
Rangitoto wharf, Jan 2014. D. nigra, O. nigricans, S. breviculus
Islington Bay, Rangitoto, May 1930**. B. quoyi, D. citrina (MA76185), E. maoria (MA78320), H. zelandiae, O. nigricans
Bird Islands, Motutapu Island, Mar 2014. B. ornata, B. quoyi, D. citrina, D. nigra, H. zelandiae, L. ophione, O. nigricans
Emu Bay, Motutapu Island, Apr 2013. D. citrina, D. nigra, O. nigricans, S. breviculus
Station Bay, Motutapu Island, Feb 2012. B. quoyi, D. citrina, O. nigricans, P. maculata
Otata Is, Noises Group, Jun 2007**. A. luctuosa (MA36464), T. rudmani
Nth end Motuihe Island, Jan 2000, Mar 2010. A. luctuosa, B. quoyi, D. citrina, L. ophione, O. nigricans, P. auriformis
(MA119218), P. maculata, S. breviculus
Ocean and West beaches, Motuihe Island, Jun 2010. M. cylindrica, P. maculata
Browns Island, Jan 2000. B. quoyi, D. citrina (MA105071), O. nigricans, S. breviculus
Crusoe Island, Sep 2009. A. luctuosa (MA118956), D. citrina, O. nigricans, P. milleri (MA118955), P. maculata, S. breviculus
Motukaha Island, Waiheke Is, Aug 2009. Aphleodoris sp. (MA118998), D. citrina, O. nigricans, P. maculata (MA118947),
S. breviculus
Enclosure and W Bays, Waiheke Is, Apr 1995*, Mar 1997*. A. argus, B. leachii, B. quoyi, L. ophione, S. breviculus
Church Bay, Waiheke Is, Apr 1995*. P. maculata
Oneroa, Waiheke Is, Jan 1987*, Feb 1988*, Jul 1988*, Dec 1989*, Apr 1995*, Sep 1999*, Aug 2009*, Jan 2014*.
A. luctuosa, B. ornata, B. quoyi, D. citrina, D. granulosa, E. maoria, H. zelandiae, L. ophione, M. cylindrica (MA29498),
O. nigricans, P. anis, P. milleri, P. maculata, R. muscula, R. oruaensis, S. breviculus
Kennedy Pt, Waiheke Is, Mar 2009. B. leachii, B. quoyi, D. citrina, O. nigricans
Te Matuku Bay, Waiheke Is, Sep 1996. B. quoyi, H. zelandiae, O. nigricans (MA133334), P. maculata
Man o’War Bay, Waiheke Is, Aug 2009. B. quoyi, L. ophione, O. nigricans, P. maculata
Rotoroa Island, Oct 2007. P. angasi (MA119011), B. quoyi, O. nigricans
Herald Is, Waitemata Harbour, Mar 2013. O. nigricans
Te Atatu Peninsula, Waitemata Harbour, Mar 2013. O. nigricans
Pt Chevalier, TP**. H. zelandiae
Meola Reef, Waitemata Harbour, May 1930**, Sep 1994, Jun 1996, Oct 2005. A. luctuosa (MA78278), B. ornata, B. quoyi,
D. citrina, D. krusensternii, D. nigra (MA130899), D. granulosa, J. pantherina (MA131859), O. nigricans, P. milleri,
P. maculata (MA87095), S. breviculus
Herne Bay, Nov 1994**, Feb 1996**, TP**. B. leachii (MA130530), B. quoyi, H. zelandiae (MA92033), M. cylindrica
Watchman Island, Waitemata Harbour, May 1997. B. ornata, D. citrina, O. nigricans, S. breviculus,
Fitzpatrick Bay, Waitemata Harbour, Jun 1996. A. lurana (MA131290)
Hobson Bay, Jun 1998. B. leachii, O. nigricans
Orakei Basin, Sep 1991*. B.leachii (MA75226)
Okahu Bay, Jun 1998*. B. ornata, D. citrina, O. nigricans
Mission Bay-St Heliers, May 1997. D. citrina, D. krusensternii
St Heliers-W Tamaki Pt, Oct 1960**, 2004. B. medietas (MA115126), B. ornata, D. citrina, J. pantherina, O. nigricans, S. breviculus
West Tamaki Pt, Jul 2004, Sep 2004, Mar 2006. A. lanuginata, A. lurana, B. quoyi, D. citrina, M. cylindrica, O. nigricans,
P. maculata, R. muscula
Karaka Bay-Glendowie, Aug 2002, 2003, 2004, Jul 2010 A. mollicella (MA113210), A. lanuginata (MA114105), B. leachii
(MA102567), B. medietas, B. quoyi, D. citrina, H. zelandiae, M. cylindrica (MA114102), O. nigricans, R. muscula
Tahuna Torea-Point England, Tamaki Estuary, Oct 2002, Jan 2003 B. leachii, B. quoyi, D. citrina, H. zelandiae, O. nigricans,
P. auriformis (MA112619), P. maculata, P. taronga
Upper Tamaki Estuary, Sep 2003. H. zelandiae, O. nigricans
Halfmoon Bay, Tamaki Estuary, Jun 2002. D. citrina (MA111352), H. zelandiae, O. nigricans
Panmure Bridge-Half Moon Bay, Oct 1986*, Sep 1988*, Jul 2002, Jun 2003 D. citrina, H. zelandiae, M. cylindrica,
O. nigricans, P. auriformis, P. taronga
Little Bucklands-Bucklands Beach, Apr 2003. B. ornata, B. leachii, B. quoyi, D. citrina, D. nigra, E. maoria, H. zelandiae,
M. cylindrica, O. nigricans, P. milleri, P. maculata, R. muscula
88
Bucklands Beach-Musick Pt, Feb 1995*, Jul 2000*, Apr 2003, Oct 2003, Mar 2006, Aug 2010. P. auriformis (MA119275).
A. lanuginata, B. leachii, B. quoyi, D. citrina, D. nigra, E. maoria (MA92806), H. zelandiae, M. cylindrica, O. nigricans,
P. auriformis, P. maculata (MA102566), P. taronga, R. muscula
Musick Pt-Eastern Beach, Aug 2004, Sep 2006, Sep 2007*, Dec 2007*, Aug 2010, Apr 2011 A. mollicella, B. ornata, B. quoyi, D.
citrina, D. krusensternii (MA119595), D. nigra (MA118313), L. ophione, O. nigricans, R. muscula, T. bractea (MA118155)
Eastern Beach Sth end, Jun 2014. A. mollicella, B. leachi, D. krusensternii, D. nigra, M. cylindrica, O. nigricans
Mellons Bay, Feb 2005, 2010**. A. mollicella, B. leachii, B. quoyi, D. krusensternii, D. nigra, H. zelandiae, O. nigricans, S.breviculus
Howick Beach, May 1954**, Nov 1999, Jan 2000, Oct 2006, Jul, Aug 2012. B. leachii (MA142020), B. quoyi, D. densioni
(MA117928), D. nigra, H. zelandiae, L. ophione (MA104746), O. nigricans, P. milleri, R. muscula
Howick Beach-Cockle Bay, Aug 2011. B. leachii D. krusensternii, D. nigra, H. zelandiae, M. cylindrica, O. nigricans, R. muscula
Mangemangeroa Estuary, **. P. dendritica (MA120714)
Motukaraka Island, Beachlands, Sep 1997, Feb 2014, B. quoyi, D. citrina, D. nigra, H. zelandiae, L. ophione, M. cylindrica,
O. nigricans, P. maculata, R. muscula, S. breviculus
Omana, Mar 2003, 2004*. B. leachii (MA115213), B. quoyi, D. citrina (MA115214), H. zelandiae
Maraetai Pt, Mar 2001. B. leachii, L. ophione, R. muscula
Maraetai, Magazine Pt, Aug 2014. B. quoyi, D. citrina, D. krusensternii, L. ophione, O. nigricans, R. muscula
Nth side Duders Regional Park, May 2003. B. leachii, B. quoyi, D. nigra, L. ophione, O. nigricans, R. muscular
Mataitai Bay, Mar 2015. D. citrina
Pouto Pt, Mar 2007. O. nigricans
West end Kawakawa Bay, Mar 2006. B. quoyi, D. citrina, O. nigricans
East end Kawakawa Bay, Jun 2004. D. citrina, H. zelandiae, O. nigricans, P. maculata
Nth Orere Pt, Apr 2015. D. krusensternii
Sth Orere Pt, May 2005. B. quoyi, D. citrina, R. muscula, O. nigricans
Matingarahi Pt, Firth of Thames, Jun 2007. O. nigricans
Great Barrier Island and Coromandel Peninsula
Whangapoua, Great Barrier Is, Jun 2000, Mar 2002, TP**. A. lanuginata (MA111094), A. parvula (MA111092), B. citrina
(MA111093), B. lineata, B. quoyii (MA111635), C. thetidis, H. zelandiae (MA103110), S. breviculus (MA110813)
Rakitu Is, Great Barrier, Jan 1981*, TP**. P. anis
Katherine Bay-Kawa, Great Barrier Is, Mar 2002. B. citrina (MA110228), B. leachii, B. quoyi, C. amoenum, O. nigricans,
R. oruaensis, S. breviculus
Port Abercrombie, Great Barrier Is, Jan 1991*, Dec 1992, Mar 2002. B. vernicosa, B. citrina, B. leachii, B. quoyi,
C. amoenum, L. ophione
Fitzroy, Great Barrier Is, Dec 1993. A. luctuosa (MA80958), G. aureomarginatus, P. angasi, P. auriformis
Kaikoura Is, Great Barrier Is, 1995*, Dec 2009. D. brazieri, D. citrina, D. krusensternii, O. nigricans, S. breviculus
Oneura Bay, Great Barrier Is, Mar 2002. B. ornata, B. leachii (MA1100912), B. quoyi, C. amoenum, D. nigra, D. granulosa
(MA110922), H. zelandiae, O. nigricans, R. muscula
Rangiahua Is, Great Barrier. Jan 1985*, 1991*. C. thetidis, C. willani, P. anis
Blind Bay, Great Barrier, Dec 1986*. A. argus
Schooner Bay, Great Barrier Is, Nov 2005. A. luctuosa, C. thetidis, D. nigra, R. muscula
Tryphena, Great Barrier Is, Feb 1991*, TP**. A. lanuginata, B. ornata, B. quoyi, C. amoenum, C. thetidis, D. brazieri,
D. nigra, D. wellingtonensis, H. zelandiae, L. ophione, M. cylindrica, O. nigricans, R. oruaensis, S. breviculus
Medlands, Great Barrier Is, Dec 1981*, Feb 2004. A. luctuosa (MA114993), A. parvula, B. lineata, B. ornata (MA114999),
B. quoyi, L. ophione, O. nigricans (MA114997), R. oruaensis (MA116696), S. breviculus
Cuvier Island, TP**. C. thetidis
Whangapoua-Mahurangi, east Coromandel Peninsula, Jun 2000. A. keraudreni, O. nigricans, S. breviculus (MA103156)
Kuaotunu, east Coromandel Peninsula, Mar 1946**. A. parvula
Whitianga, east Coromandel Peninsula, 2010**. P. maculata
Hahei, east Coromandel Peninsula, Dec 2000**. A. luctuosa (MA116991), C. rubiginosa
Hot Water Beach, TP**. P. anis
Pauanui, east Coromandel Peninsula, Nov 2007*. A. lanuginata (MA30167)
Opoutere, east Coromandel Peninsula, 2005**. F. pinnata (MA118314), M. cylindrica, R. katipoides
Fletchers Bay, Coromandel Peninsula, Dec 2013. B. ornata, B. citrina
Waitete Bay, west Coromandel Peninsula, Jan 2013. B. citrina, O. nigricans
Wyuna Bay-Little Passage, west Coromandel Peninsula, Jan 2013. B. leachii, B. quoyi, O. nigricans, P. angasi, P. auriformis
(MA120174)
Wilson Bay, west Coromandel Peninsula, Dec 2013. D. citrina, O. nigricans
Te Mata, west Coromandel Peninsula, Oct 2014. L. ophione, O. nigricans
Margaret S. Morley & Bruce W. Hayward
89Intertidal records of ‘sea slugs’
Taxon
Biogeography
Kermadecian
Aupourian
Cookian
Forsterian
Moriorian
Antipodean
Animal colour
Shell
Figure
Acteonidae
Pupa anis (Hutton, 1873) Cos A C Creamy white Ext 1
Bullinidae
Bullina lineata (Gray, 1825) IP K A Cream with blue margin Ext 1
Aplustridae
Hydatina zonata (Lightfoot, 1786) IP AExt 1
Hydatina physis (Linnaeus, 1758) Cos K A Pink Ext 1
Cephalaspidea: Cylichnidae
Cylichna thetidis Hedley, 1903 Aus K A C Ext 2
Philinidae
Philine angasi (Crosse
and Fischer, 1865) Aus K A C White Int 2
Philine auriformis Suter, 1900 E A C F White Int 2
Retusidae
Retusa oruaensis (Webster, 1908) E A C F M Creamy white 2
Aglajidae
Melanochlamys cylindrica
Cheeseman, 1881 E A C F An Black Int 3
Melanochlamys lorrainae
(Rudman, 1968) E A C White to grey Int 3
Philinopsis taronga Allan, 1933 Aus ABrown with yellow spots Int 3
Runcinidae
Pseudoilbia zelandica Miller
and Rudman, 1968 E A Black longitudinal bands,
white front and back Int 3
Runcina katipoides (Miller
and Rudman, 1968) E A Dk grey, with red
central stripe Int 3
Runcinella zelandica Odhner, 1924 E A Dk green with
irregular pink bands
and white splotches
Int 3
Bullidae
Bulla vernicosa (Pilsbry, 1893) SP K A Ext 4
Bulla quoyii Gray, 1843 Aus A C Ext 4
APPENDIX 2: Summary of distribution and features of intertidal and shallow subtidal sea slugs of north-
ern New Zealand. The known New Zealand-wide distribution of each species is given in
terms of New Zealand biogeographic provinces of Powell (1955) and listed in Spencer et
al. (2014). E = Endemic to New Zealand. Biogeography: Aus = Australasian; Cos = cosmopolitan in
warm waters; E = NZ endemic; IP = Indo-Pacific;Pac = Pacific; SP = South Pacific; Shell: Ext = easily seen
externally; Int = internal and not seen externally
90
Taxon
Biogeography
Kermadecian
Aupourian
Cookian
Forsterian
Moriorian
Antipodean
Animal colour
Shell
Figure
Haminoeidae
Haminoea zelandiae (Gray, 1843) E A C Dark grey Ext 5
Aplysiomorpha: Aplysiidae
Aplysia argus (Ruppell
and Leuckart, 1830) Cos K A Olive green, with
black reticulate lines Int 6
Aplysia juliana (Quoy
and Gaimard, 1832). Cos A C F Choc brown to black,
with white spots Int 6
Aplysia keraudreni Rang, 1828 SP A C F Kelp brown with white
mottles, black netted lines Int 6
Aplysia parvula Guilding, 1863 Cos A C F Kelp brown with
white dots Int 6
Bursatella leachii Blainville, 1817 Pac A C Brownish-green, black
speckles, emerald
green patches
No 7
Dolabrifera brazieri
G.B. Sowerby, 1870 Aus K A Green marbled brown,
white and pink Int 7
Stylocheilus longicauda
Quoy and Gaimard, 1832 Cos AGreen with brown
lines and spots
ringed with black
No 7
Sacoglossa: Limapontiidae
Ercolania felina (Hutton, 1882) E A C F Grey-black, central
white stripe No 8
Stiliger smaragdinus Baba, 1949 Pac ABright green No 8
Placida dendritica (Alder
and Hancock, 1843) Cos A C F An Green No 8
Placobranchidae
Elysia maoria Powell, 1937 Aus A C Dark green No 8
Elysia n.sp. Aus APale green with
blue spots & bright
blue-green margin
No 8
Pleurobranchomorpha:
Pleurobranchidae
Berthella medietas Burn, 1962 Aus A C F M An Orange to bronze Int 9
Berthella ornata (Cheeseman, 1878) E A C F M Cream-white with dark
red-brown blotches Int 9
Berthellina citrina (Ruppell
and Leuckart, 1828) IP A C Orange to pale
yellow or red Int 9
Pleurobranchaea maculata
(Quoy and Gaimard, 1832) IP A C F Grey to black with
dark wavy lines No 10
Nudibranchia: Cadlinidae
Cadlina
willani Miller, 1980 E A C White with central
yellow stripe and border No 11
Margaret S. Morley & Bruce W. Hayward
91Intertidal records of ‘sea slugs’
Taxon
Biogeography
Kermadecian
Aupourian
Cookian
Forsterian
Moriorian
Antipodean
Animal colour
Shell
Figure
Chromodorididae
Ceratosoma amoenum
(Cheeseman, 1886) Aus A C White with orange
patches and purple
gills and rhinophores
No 11
Goniobranchus aureomarginatus
(Cheeseman, 1881) E A C F M White bordered
with gold line No 11
Discodorididae
Alloiodoris lanuginata
(Abraham, 1877) E A C F M Red-brown to grey
with brown and white
spots; green gills
No 12
Atagema molesta (Miller, 1989) E A Fawn brown No 12
Hoplodoris nodulosa (Angas, 1864) E A C F Yellow to grey with
brown blotches, gills
and rhinophores
No 12
Jorunna pantherina (Angas, 1864) Aus APeachy, yellow to black
with marbled ring patches No 12
Rostanga muscula (Abraham, 1877) E A C Tomato red with
dark spots No 12
Dorididae
Aphelodoris luctuosa
(Cheeseman, 1882) E A C F M Cream with brown
stripes, blotches and
concentric lines
No 13
Aphelodoris sp. E C Cream with tan blotches No 13
Doris granulosa (Pease, 1860) Aus A C F M Yellow No 14
Doris wellingtonensis
Abraham, 1877 Aus K A C F M Khaki brown to yellow No 14
Dendrodoridae
Dendrodoris citrina
(Cheeseman, 1881) E A C F Lemon yellow to
orange with numerous
small white dots
No 15
Dendrodoris krusensternii
(Gray, 1850) IP ALight brown and
peacock blue with
concentric brown lines
No 16
Dendrodoris nigra (Stimpson, 1855) IP A C Black with white
tipped rhinophores No 16
Polyceridae
Plocamopherus imperialis
Angas, 1864 IP AScarlet with sm dk spots,
2 cream stripes; white
tipped rhinophores
No 17
Polycera hedgpethi Er. Marcus, 1964 Cos A C Cream peppered
with black, white
and gold papillae
No 17
Polycera melanosticta Miller, 1996 Aus ATranslucent white
to greyish-brown
with black spots
No 17
92
Taxon
Biogeography
Kermadecian
Aupourian
Cookian
Forsterian
Moriorian
Antipodean
Animal colour
Shell
Figure
Thecacera pennigera
(Montagu, 1815) Cos AWhite with orange
and black spots No 17
Proctonotidae
Caldukia rubiginosa Miller, 1970 E A Reddish-brown; yellow
and iridescent blue cerata No 17
Janolus novozealandicus
(Eliot, 1907) E A C F Translucent with brown
& white speckles;
diverticula blue & orange
No 17
Vassieridae
Vayssierea cinnabarea Ralph, 1944 E A C Dull red No 17
Fionidae
Fiona pinnata (Eschscholtz, 1831) Cos A C F M An Cream or bu with
orange patch No 17
Onchidorididae
Acanthodoris mollicella
Abraham, 1877 E A C F An Mottled black, brown
and ochre-yellow No 18
Goniodorididae
Goniodoris n.sp. E A Unknown No 18
Okenia plana Baba, 1960 Pac ALilac and red-brown
spotted No 18
Trapania rudmani Miller, 1981 E A C White with yellow stripes No 18
Dotidae
Doto pita Er. Marcus, 1955 Cos A C Translucent white
with brown, yellow
and lilac spots
No 19
Arminidae
Dermatobranchus rubidus
(Gould, 1852) Aus ADark orange and white No 19
Tritoniidae
Tritonia incerta Bergh, 1904 E A C Rose-pink, apricot or red,
with white gills and foot No 19
Flabellinidae
Flabellina albomarginata
(Miller, 1971) E A C F White, with red cerata No 19
Tularia bractea (Burn, 1962) Aus A C Transparent white-yellow-
brown with white ecks,
red-greenish diverticula
No 19
Eubranchidae
Eubranchus agrius Er. Marcus, 1959 Pac A C F Transparent with sm
patches and stripes
of yellow to red
No 19
Margaret S. Morley & Bruce W. Hayward
93Intertidal records of ‘sea slugs’
Taxon
Biogeography
Kermadecian
Aupourian
Cookian
Forsterian
Moriorian
Antipodean
Animal colour
Shell
Figure
Tergipedidae
Trinchesia beta (Baba
and Abe, 1964) Pac A C Transparent speckled
purple, gold band,
pink diverticula
No 19
Trinchesia reexa (Miller, 1977) E C F Transparent to white,
fawn speckled diverticula No 19
Aeolidiidae
Aeolidiella drusilla Bergh, 1900 Aus A C F M Bu with white
blotches with brown
and orange cerata
No 20
Anteaeolidiella lurana (Bergh, 1888) Cos AOrange to brown,
with orange stripes
and white blotches
No 20
Baeolidia australis (Rudman, 1982) Aus ABrown with white
spots. Brown, crimson
and gold cerata
No 20
Burnaia helicochorda (Miller, 1988) Aus ATransparent orange-
brown, white-
tipped cerata
No 20
Facelinidae
Phidiana milleri Rudman, 1980 E A C Translucent with orange
head, tentacles and cerata No 20
Babakinidae
Babakina caprinsulensis
(Miller, 1974) E A Pale mauve with
white, yellow and
orange features
No 21
Glaucinidae
Glaucus atlanticus Forster, 1777 Cos K A C Glowing blue,
silver and purple No 21
Umbraculidae
Umbraculum umbraculum
(Lightfoot, 1786) Cos K A C Grey to yellow,
brown or red Ext 21
Velutinidae
Lamellaria ophione Gray, 1850 E K A C Yellow, orange,
grey or white Int 22
Onchiidae
Onchidella nigricans (Quoy
& Gaimard, 1835) Aus A C F M Black-grey No 23
Fissurellidae
Scutus breviculus (Blainville, 1817) E A C F Black Int 24
... The species P. maugeansis is restricted to southeastern Australia, where it occurs intertidally in tide pools on rocky platforms and subtidally on muddy seagrass beds (Burn 1966a(Burn , 1969Hales 2010;Burn and Wilson 2011). Papawera zelandiae is endemic to New Zealand, where it is relatively common on intertidal and subtidal sheltered sandy-mud flats and on rocky shores with algae turfs (Rudman 1971a(Rudman , 1971bPowell 1979;Morley and Hayward 2015). ...
... Rudman (1971aRudman ( , 1971b first described several anatomical features and its diet. Willan (1979), Hayward (1979) and Morley and Hayward (2015) also provided data on diet as well as ecological aspects such as habitat and distribution. Burn (1966a) described the species Haminoea maugeansis from Victoria, South Australia, and Tasmania, based on the shell, external morphology, and several anatomical features of the digestive system (jaws, radula and gizzard plates) but only included a drawing of the shell and one row of the radula (Burn 1966a, p. 330, Figures 1, 2). ...
... The species is most frequently recorded around New Zealand's North Island and the northern parts of the South Island (Rudman 1971a;Rudman 1971bRudman , 1999Rudman , 2003Rudman , 2006Powell 1979;Furneaux 2003;Morley and Hayward 2015). However, the species also can be found on the southern coast of the South Island (based on one shell housed at the AMS c.457260), but to our knowledge, it has not been observed alive so far south. ...
Article
Full-text available
The genus Papawera includes two species of haminoeid snails found only in temperate waters of New Zealand and southeastern Australia. In this work, we redescribe the Papawera species based on characters of their external morphology, shells, and anatomical features such as radulae, jaws, gizzard plates, and male reproductive systems, using for the first time, scanning electron microscopy. A multi-locus phylogenetic hypothesis and the species delimitation method Automatic Barcode Gap Discovery based on DNA sequences of the cytochrome c oxidase subunit I gene were used to corroborate species status. The type species of the genus, P. zelandiae, is restricted to New Zealand and P. maugeansis is only known from South Australia, Victoria, and Tasmania. These species are easily distinguished externally by the shape of the shell, colouration of the living animal, and morphology of the cephalic shield. Anatomically they have differences in the number of marginal teeth, distribution of rods in the gizzard plates, and anatomy of the fundus in the male reproductive system.
... northern New Zealand (Rudman, 2003), including the Kermadec Islands (Morley & Hayward, 2015). According to Morley & Hayward (2015), the presence of this species in New Zealand is the result of a humanmediated introduction. ...
... northern New Zealand (Rudman, 2003), including the Kermadec Islands (Morley & Hayward, 2015). According to Morley & Hayward (2015), the presence of this species in New Zealand is the result of a humanmediated introduction. ...
Article
Molecular and morphological data from newly collected specimens and a review of the literature and type material indicate that the widespread tropical sea hare Dolabrifera dolabrifera is a species complex of five genetically distinct taxa. The name Dolabrifera dolabrifera is retained for a widespread species in the Indo-Pacific tropics. Dolabrifera nicaraguana is endemic to the eastern Pacific. Dolabrifera ascifera, D. virens and a new species described herein are restricted to the tropical Atlantic, with partially overlapping ranges in the Caribbean region and St. Helena. The temperate Pacific species Dolabrifera brazieri is also distinct and endemic to temperate southeastern Australia and New Zealand. These species of Dolabrifera constitute highly divergent lineages and most contain unique internal anatomical characteristics, particularly in the male reproductive morphology and shell shape, making them relatively easy to identify upon dissection. However, externally all these species are extremely variable in colour pattern and morphology and are virtually indistinguishable. This is particularly problematic for identification in the Atlantic Ocean where three species co-occur in the Caribbean region.
... northern New Zealand (Rudman, 2003), including the Kermadec Islands (Morley & Hayward, 2015). According to Morley & Hayward (2015), the presence of this species in New Zealand is the result of a humanmediated introduction. ...
... northern New Zealand (Rudman, 2003), including the Kermadec Islands (Morley & Hayward, 2015). According to Morley & Hayward (2015), the presence of this species in New Zealand is the result of a humanmediated introduction. ...
Article
Full-text available
Molecular and morphological data from newly collected specimens and a review of the literature and type material indicate that the widespread tropical sea hare Dolabrifera dolabrifera is a species complex of five genetically distinct taxa. The name Dolabrifera dolabrifera is retained for a widespread species in the Indo-Pacific tropics. Dolabrifera nicaraguana is endemic to the eastern Pacific. Dolabrifera ascifera, D. virens and a new species described herein are restricted to the tropical Atlantic, with partially overlapping ranges in the Caribbean region and St. Helena. The temperate Pacific species Dolabrifera brazieri is also distinct and endemic to temperate southeastern Australia and New Zealand. These species of Dolabrifera constitute highly divergent lineages and most contain unique internal anatomical characteristics, particularly in the male reproductive morphology and shell shape, making them relatively easy to identify upon dissection. However, externally all these species are extremely variable in colour pattern and morphology and are virtually indistinguishable. This is particularly problematic for identification in the Atlantic Ocean where three species co-occur in the Caribbean region.
... Nudibranchs from the Genus Plocamopherus are known to be able to flatten their tail into a paddle-like shape, where it could use its tail for short locomotion (Morley and Hayward 2015). Inhabit intertidal rocky shores (Nimbs and Smith 2017;Gosliner et al. 2019). ...
Article
Full-text available
Andrimida A. 2022. New records of nudibranchs and a sacoglossan (Gastropoda: Heterobranchia) from Sempu Strait, Indonesia. Indo Pac J Ocean Life 6: 1-9. This study reports the additional records of fourteen heterobranch sea slug species from Sempu Strait, East Java, Indonesia, in which thirteen species belong to the nudibranchia order, and one species belongs to the sacoglossa order. These species belong to seven families: Plakobranchidae (1 species), Polyceridae (1 species), Discodorididae (2 species), Chromodorididae (7 species), Flabellinidae (1 species), Eubranchidae (1 species), and Facelinidae (1 species). All of these species are recorded for the first time from the Sempu Strait. There are two species that only could be identified to its genus, which are Chromodoris sp. and Eubranchus sp. This study also found additional record of Plocamopherus imperialis Angas, 1864 on the tropical waters, where this species formerly believed only exclusively distributed along the subtropical waters of Eastern Australia and Northern New Zealand. Combined with the previous study on Sempu Strait, to date, 59 species of heterobranch sea slug have been reported and recorded in this area. Species identification was conducted by carefully examining the morphological features of each specimens encountered during the surveys.
Article
Full-text available
An annotated checklist of sea slugs from Gujarat coastal waters was prepared, based on published literature and field observations from 2014–2019. Ninety-five species from 62 genera belonging to 29 families were recorded. Species are listed with synonyms and distribution status.
Article
Full-text available
Four-hundred and fifty-nine species of marine biota (154 gastropods, 74 bivalves, 42 seaweeds, 40 foraminifera, 34 ostracods, 25 crabs and shrimps, 16 amphipods, 10 echinoderms, 10 chitons, 9 isopods, 9 polychaetes, 7 barnacles and smaller numbers of other groups) are recorded from a 4 km long stretch of coast around Whangapoua Beach, north-east Coromandel Peninsula, including New Chums Beach and inside the entrance to Whangapoua Harbour.
Article
Full-text available
The origin of tetrodotoxin (TTX) is highly debated; researchers have postulated either an endogenous or exogenous source with the host accumulating TTX symbiotically or via food chain transmission. The aim of this study was to determine whether the grey side-gilled sea slug (Pleurobranchaea maculata) could obtain TTX from a dietary source, and to attempt to identify this source through environmental surveys. Eighteen non-toxic P. maculata were maintained in aquariums and twelve were fed a TTX-containing diet. Three P. maculata were harvested after 1 h, 24 h, 17 days and 39 days and TTX concentrations in their stomach, gonad, mantle and remaining tissue/fluids determined using liquid chromatography-mass spectrometry. Tetrodotoxin was detected in all organs/tissue after 1 h with an average uptake of 32%. This decreased throughout the experiment (21%, 15% and 9%, respectively). Benthic surveys at sites with dense populations of toxic P. maculata detected very low or no TTX in other organisms. This study demonstrates that P. maculata can accumulate TTX through their diet. However, based on the absence of an identifiable TTX source in the environment, in concert with the extremely high TTX concentrations and short life spans of P. maculata, it is unlikely to be the sole TTX source for this species.
Article
Full-text available
Open-coast and deep-water ecosystems have been less disrupted by invasion than estuaries, but a notable exception is the establishment of multiple species of the sea slug genus Philine in the northeastern Pacific. These large slugs spread from San Francisco Bay to the whole of the US west coast in 5 years during the 1990s, and are abundant from intertidal mudflats to soft-sediment bottoms >300 m deep along the open coast. Voracious predators that secrete acid, Philine spp. have few natural enemies and substantial impacts on native bivalve communities. Despite their ecological significance, the identity and number of invasive Philine spp. in the US has remained controversial. Here, we adopt a molecular approach to identify the species commonly found along the US west coast. We compared mitochondrial 16S gene sequences of reference specimens from the native range of seven possible invaders against sequences of 66 specimens collected from southern California to Oregon from intertidal and subtidal habitats. All slugs from southern California and Oregon were confirmed as P. auriformis, and most shared haplotypes with samples from New Zealand. Larger slugs from San Francisco, Tomales Bay and Bodega Harbor, believed to represent 3–4 species, were all identified as P. orientalis. Molecular data support a recent morphological analysis that erected two proposed species, P. paucipapillata from Hong Kong and P. quadripartita from Europe, which are genetically distinct from anatomically similar species. For taxonomically challenging groups like sea slugs, genetic data can illuminate cryptic invasions and provide a backdrop for further studies of the ecological impacts of introduced marine species.
Article
Full-text available
Seventy new records of Mollusca (7 chitons, 41 gastropods, 22 bivalves) are added to the existing species list for the Kawerua area, on the west coast of North Auckland, bringing the total to 192 species (13 chitons, 125 gastropods, 51 bivalves, 3 cephalopods). These Kawerua records provide geographic range extensions for 21 species of mollusc.