Opisthobranchia

Transcription

Opisthobranchia

Undergraduate thesis in biology
Opisthobranchia: A taxonomic and biological review
with emphasis on the families Chromodorididae and
Phyllidiidae together with field notes from South East
Sulawesi, Indonesia
Lars Karlsson
LiU-IFM-Biol-Ex-892
Avdelning, Institution
Division, Department
Datum
Date
Avdelningen för biologi
010330
Institutionen för Fysik och mätteknik
Språk
Language
Engelska/English
Rapporttyp
Report category
x Examensarbete
x D-uppsats
ISBN
LiU-IFM-Biol-Ex-892
ISRN
LiU-Biol-Ex-350
Handledare: Per Milberg
Institution: Biology Department, IFM, University of Linköping
URL för elektronisk version
Titel
Title: Opisthobranchia: A taxonomic and biological review with emphasis on the families Chromodorididae and Phyllidiidae
together with field notes from South East Sulawesi, Indonesia
Författare
Author: Lars Karlsson
Sammanfattning
Abstract
This study gives an overview regarding Opisthobranchia (Mollusca, Gastropoda) and its taxonomy and some important biological
facts. It also contains two parts of information regarding fieldwork conducted in Indonesia (Sulawesi) in cooperation with
“Operation Wallacea” (a joint venture between a British company and an Indonesian research institute).
The first of the two parts regarding fieldwork, gives a checklist of “Opisthobranchs of the Wakatobi Marine National Park”
containing a total of 297 identified or suspected (provisionally identified) species (1997-1999). In the survey season 1999: 3523
opisthobranch observations were recorded, belonging to 235 species (identified or suspected) and 73 identified genera. During the
whole survey 60 % of the opisthobranchs were identified to species level. The largest order was the Doridina, both in species
number and number of genera. Fourteen percent of the opisthobranch species found in 1999 made up seventy-five percent of the
total number of observations recorded.
The second of the two parts regarding fieldwork is a description and analysis of spatial distribution of nudibranchs according to
benthic structure (especially the families: Chromodorididae and Phyllidiidae since they are the most abundant). This survey was
conducted at two sites, at two depths (around 3 and 12 m) and at two times a day (“am/pm”). During this survey 252 nudibranchs
belonging to at least 19 species were recorded. According to a Detrended Correpondance Analysis, species composition seems to
vary with benthic forms and depth.
Nyckelord
Keywords: Opisthobranchia, Nudibranchia, Chromodorididae, Phyllidiidae, Operation Wallacea, Sulawesi, opisthobranch
surveys, spatial distribution
Contents
1
2
3
4
5
Contents
Abstract
Review of opisthobranch taxonomy & biology
1.1 Introduction
1.2 Taxonomy of Gastropoda/Opisthobranchia
1.3 Morphology, ecology and physiology in Opisthobranchia
1.4 Nudibranchia
1.5 Chromodorididae
1.6 Phyllidiidae
1.7 References
Checklist of opisthobranchs of the Wakatobi National Park,
SE Sulawesi, Indonesia
2.1 Introduction
2.2 Methods
2.3 Results
2.4 Discussion
2.5 References
Spatial distribution of nudibranchs according to benthic structure
3.1 Introduction
3.2 Material and methods
3.3 Results
3.4 Discussion
3.5 References
Conclusions and recommendations
Acknowledgements
Plates
Appendices
1
1
2
3
3
3
6
11
15
18
18
21
21
21
23
26
28
29
29
29
31
35
36
37
38
A
B
I
II
Opisthobranchia: A taxonomic and biological review with emphasis on
the families Chromodorididae and Phyllidiidae together with field
notes from South East Sulawesi, Indonesia
Lars Karlsson, Biology Department, IFM, University of Linköping, 2001.
Abstract
This study gives an overview regarding Opisthobranchia (Mollusca, Gastropoda) and its
taxonomy and some important biological facts. It also contains two parts of information
regarding fieldwork conducted in Indonesia (Sulawesi) in cooperation with “Operation
Wallacea” (a joint venture between a British company and an Indonesian research institute).
The first of the two parts regarding fieldwork, gives a checklist of “Opisthobranchs of the
Wakatobi Marine National Park” containing a total of 297 identified or suspected
(provisionally identified) species (1997-1999). In the survey season 1999: 3523 opisthobranch
observations were recorded, belonging to 235 species (identified or suspected) and 73
identified genera. During the whole survey 60 % of the opisthobranchs were identified to
species level. The largest order was the Doridina, both in species number and number of
genera. Fourteen percent of the opisthobranch species found in 1999 made up seventy-five
percent of the total number of observations recorded.
The second of the two parts regarding fieldwork is a description and analysis of spatial
distribution of nudibranchs according to benthic structure (especially the families:
Chromodorididae and Phyllidiidae since they are the most abundant). This survey was
conducted at two sites, at two depths (around 3 and 12 m) and at two times a day (“am/pm”).
During this survey 252 nudibranchs belonging to at least 19 species were recorded. According
to a Detrended Correpondance Analysis, species composition seems to vary with benthic forms
and depth.
2
1 Review of opisthobranch taxonomy & biology
1.1 Introduction
Opisthobranchs or “sea slugs” are a large, exclusively (except for the freshwater Ancylodoris
baicalensis) marine group within the Gastropoda (Wägele & Willan, 2000). They are often
colourful and display several forms of crypsis, special resemblance, aposematic coloration and
mimicry (Gosliner & Behrens, 1990; Tullrot, 1998). Most opisthobranchs lack the protection of
a shell as adults (some of them do have a shell but it is not functioning as a protective device
since it is reduced and sometimes also internal). Instead these organisms rely on other
defences. Some opisthobranchs use spicules as an alternative to shell, others are cryptic, some
utilize the nematocysts from their prey and a few use autotomy. One common defensive
systems, though, is chemical defence (Faulkner & Ireland, 1977; Todd, 1981; Faulkner &
Ghiselin, 1983; Gunthorpe & Cameron, 1987; Avila, 1995; Cimino et al., 1999). This is the
case for both Chromodorididae and Phyllidiidae (Nudibranchia). Both groups prey on sponges,
which often contain bad tasting or even toxic compounds. The opisthobranchs then accumulate
(or convert) some of these compounds and thereby become less interesting for potential
predators. This habit has made them interesting for pharmaceutical screening as many of the
compounds they use are suspected to have pharmaceutical value (Bohlin, 1989). Another
interesting feature is that the chemical toxicity of these species often is coupled with strong
examples of aposematism. Often, the same type of strong warning coloration is used by many
different opisthobranchs. If the opisthobranchs with the similar coloration are all unpalatable,
then it is an example of Müllerian mimicry. On the other hand, if there are palatable species
among the ones with similar coloration, then it is an example of Batesian mimicry (Ruppert &
Barnes, 1994). One interesting discovery is that there is also a flatworm species, which seems
to mimic a phyllidiid nudibranch in order to avoid predation (Newman et al., 1994).
1.2 Taxonomy of Gastropoda/Opisthobranchia
There are many different and confusing views on Gastropod systematics. In most publications,
the class Gastropoda consists of three subclasses: Prosobranchia, Pulmonata and
Opisthobranchia (Ruppert & Barnes, 1994; Hayward & Ryland, 1995; Thompson, 1976). This
class is not considered by some authors to reflect the phylogeny of gastropods and a newer
(cladistic) classification divides the Gastropoda into two clades: Eogastropoda and
Orthogastropoda. In the latter group is a clade called Apogastropoda, which contain
Heterobranchia and Caenogastropoda. The former group, in turn, consists of three clades:
Heterostropha, Opisthobranchia and Pulmonata (Ponder & Lindberg, 1997). Most authors
agree that within the Opisthobranchia there are at least eight groups, although they have been
given different taxonomic ranks. These subgroups within the Opisthobranchia are (Marshall &
Willan 1999): Cephalaspidea (Bubble shells), Anaspidea (Sea hares), Notaspidea (Sidegilled slugs), Acochlidiacea (Caddis-slugs), Sacoglossa (Sap-sucking sea slugs),
Thecosomata (Sea butterflies), Gymnosomata (Naked sea butterflies) and Nudibranchia
(Nudibranchs). Sometimes other groups are included in the Opisthobranchia, for example
Pyramidellacea (Ruppert & Barnes, 1994) Pyramidellidae (Hayward & Ryland, 1995),
Entomotaeniata (Willan & Coleman, 1984), Rhodopida (Hansson, 1998) and Rhodopemorpha
(Beesley et al., 1998). Many of these groups have previously been assigned different names
and some of these synonyms are shown in Table 1 together with examples of different views
regarding levels of taxa.
3
Table 1. Systematic review of Opisthobranchia according to different authors.
Marshall & Willan,
1999
Regnum
Phylum
Subphylum
Superclassis
Classis
Subclassis
Infraclassis
Superordo
Ordo
Rudman & Willan,
1998
(In Beesley et al.,
1998)
Animalia
Mollusca
Conchifera
Gastropoda
Opisthobranchia
Gastropoda
Opisthobranchia
Cephalaspidea
Cephalaspidea =
Bullomorpha
Ordo
Anaspidea
Ordo
Notaspidea
Ordo
Acochlidia
Ordo
Sacoglossa
Ordo
Ordo
Ordo
Ordo
Thecosomata
Gymnosomata
Anaspidea =
Aplysiomorpha
Notaspidea =
Pleurobranchomorpha
Acochlidea =
Acochlidia
Sacoglossa =
Ascoglossa =
Monostichoglossa =
Pellibranchiata
Thecosomata
Gymnosomata
Rhodopemorpha
Nudibranchia =
Thompson, 1988 **
Regnum
Phylum
Subphylum
Superclassis
Classis
Subclassis
Infraclassis
Superordo
Ordo
Ordo
Ordo
Ordo
Ordo
Hansson, 1998
Nudibranchia
Behrens, 1991 and
Gosliner, 1987
Hayward & Ryland,
1995
Ruppert &
Barnes,
1994 *
Mollusca
Animalia
Mollusca
Gastropoda
Opisthobranchia
Gastropoda
Opisthobranchia
Cephalaspidea
Cephalaspidea
Anaspidea
Aplysiacea
Notaspidea
Notaspidea
Acochlidiomorph(id)a =
Acochlidioid(e)a
Sacogloss(id)a =
Ascogloss(id)a
Acochlidiacea
Acochlidioidea
Sacoglossa
Sacoglossa
Thecosomat(id)a
Gymnosomat(id)a
Rhodopida
Nudibranchi(d)a
Willan & Coleman,
1984
Thecosomata
Gymnosomata
Pyramidellidae
Nudibranchia
Thompson, 1976
Thecosomata
Gymnosomata
Pyramidellacea
Nudibranchia
Gastropoda
Heterobranchia
Euthyneura
Opisthobranchia
Cephalaspid(e)a =
Bullomorpha =
Tectibranchida
Anaspid(e)a =
Aplysiomorphi(id)a
Anaspidea =
Mollusca
Gastropoda
Opisthobranchia
Gastropoda
Opisthobranchia
Gastropoda
Opisthobranchia
Opisthobranchia
Cephalaspidea =
Bullomorpha
Bullomorpha
Cephalaspidea
Bullomorpha
Anaspidea =
Aplysiomorpha
Anaspidea
Aplysiomorpha
Notaspidea =
Pleurobranchomorpha Notaspidea
Pleurobranchomorpha
Acochlidiacea
Acochlidiacea
Sacoglossa =
Ascoglossa
Sacoglosssa
Sacoglossa
Ordo
Thecosomata
Ordo
Gymnosomata
Ordo
Entomotaeniata
Ordo
Nudibranchia
Nudibranchia
Nudibranchia
* Note: Thecosomata and Gymnosomata are sometimes called: "pteropods".
** Note: Excluding the pteropods and the pyramidellids.
Aplysiomorpha
Pleurobranchomorpha
Acochlidiacea
Sacoglosssa
Thecosomata
Gymnosomata
Nudibranchia
Going further down in the classification, a summary of opisthobranch families is presented in
Table 2. (This classification is also used by a very valuable database on the Internet, “The Sea
Slug Forum” (Rudman, 2000)).
4
Table 2. Summary of ophistobranch families (after Beesley et al, 1998).
OPISTHOBRANCHIA
Order CEPHALASPIDEA
Superfamily ACTEONOIDEA
Family Acteonidae
Family BuIlinidae
Family Hydatinidae
Superfamily RINGICULOIDEA
Family Ringiculidae
Superfamily CYLINDROBULLOIDEA
Family Cylindrobullidae
Superfamily DIAPHANOIDEA
Family Diaphanidae
Superfamily PHILINOIDEA
Family Cylichnidae
Family Retusidae
Family Philinidae
Family Philinoglossidae
Family Aglajidae
Family Gastropteridae
Superfamily HAMINOEOIDEA
Family Haminoeidae
Family Bullactidae
Family Smaragdinellidae
Family Incertae sedis
Superfamily BULLOIDEA
Family Bullidae
Superfamily RUNCINOIDEA
Family Runcinidae
Family Ilbiidae
Superfamily INCERTAE SEDlS
Family Notodiaphanidae
Order ACOCHLIDEA
Superfamily ACOCHLIDIOIDEA
Family Acochlidiidae
Family Hedylopsidae
Superfamily MICROHEDYLOIDEA
Family Asperspinidae
Family Microhedylidae
Family Ganatidae
Order RHODOPEMORPHA
Family Rhodopidae
Order SACOGLOSSA
Superfamily OXYNOOIDEA
Family Volvatellidae
Family Oxynoidae
Family Juliidae
Superfamily ELYSIOIDEA
Family Plakobranchidae
Family Elysiidae
Family Boselliidae
Family GascoigneIlidae
Family Platyhedylidae
Superfamily LIMAPONTIOIDEA
Family Caliphyllidae
Family Costasiellidae
Family Hermaeidae
Family Limapontiidae
Order ANASPIDEA
Order NUDIBRANCHA
Suborder DORIDINA
Superfamily ANADORIDOIDEA
Family Corambidae
Family Goniodorididae
Family Onchidorididae
Family Polyceridae
Family Gymnodorididae
Family Aegiretidae
Family Vayssiereidae
Superfamily EUDORIDOIDEA
Family Hexabranchidae
Family Dorididae
Family Chromodorididae
Family Dendrodorididae
Family Phyllidiidae
Suborder DENDRONOTINA
Superfamily AKEROIDEA
Family Akeridae
Superfamily APLYSIOIDEA
Family Aplysiidae
Order NOTASPIDEA
Superfamily TYLODINOIDEA
Family Tylodinidae
Family Umbraculidae
Superfamily PLEUROBRANCHOIDEA
Family Pleurobranchidae
Order THECOSOMATA
Family Limacinidae
Family Cavoliniidae
Family Peraclididae
Family Cymbuliidae
Family Desmopteridae
Order GYMNOSOMATA
Suborder GYMNOSOMATA
Family Pneumodermatidae
Family Notobranchaeidae
Family Cliopsidae
Family Clionidae
Suborder GYMNOPTERA
Family Hydromylidae
5
Family Tritoniidae
Family Bornellidae
Family Marianinidae
Family Hancockiidae
Family Dotidae
Family Scyllaeidae
Family Tethydidae
Family Phylliroidae
Family Lomanotidae
Suborder ARMININA
Family Arminidae
Family Doridomorphidae
Family Charcotiidae
Family MadreIlidae
Family Zephyrinidae
Family Pinufiidae
Suborder AEOLIDINA
Family Flabellinidae
Family Eubranchidae
Family Aeolidiidae
Family Glaucidae
Family Embletoniidae
Family Tergipedidae
Family Fionidae
1.3 Morphology, ecology and physiology in Opisthobranchia
Figure 1 shows a broad characterisation of grades of organisation in the opisthobranchs. The
circle “1” is considered to represent the oldest and most “primitive” level.
Figure 1. Grades (time-development level) of organisation in the opisthobranchs (Beesley et
al, 1998) (Reproduced by special permission from Richard Willan and Australian Biological
Resources Study, Environment Australia)
As can be seen, within each order there are usually several grades of organisation. In spite of
this, it is also possible to see that the different groups of nudibranchs are considered to have
reached the “highest” forms of organisation (Beesley et al., 1998). There is a strong coupling
between organisation grades and morphological appearances, as for instance, when the shell is
reduced or lacking completely. The first way of identifying opisthobranch orders is usually by
external examination as the morphology often gives clues to which order a specimen can be
placed. Figure 2-9 shows some typical members of eight opisthobranch groups.
a
b
Figure 2. (a) Cephalaspidea, Bulla ampulla.
(Thompson, 1976).
(b) Cephalaspidea, Chelidonura sandrana
(Thompson, 1976).
6
Figure 3. Anaspidea, Aplysia depilans
(Thompson, 1976).
Figure 4. Notaspidea, Pleurobranchus
membranaceus (Thompson, 1976).
a
b
Figure 5. (a) Acochlidiacea, Hedylopsis loricata
(Thompson, 1976).
a
(b) Acochlidiacea, Microhedyle sp
(Thompson, 1976).
b
Figure 6. (a) Sacoglossa, Stiliger fuscatus
(Thompson, 1976).
(b) Sacoglossa, Elysia marginata
(Thompson, 1976).
7
a
Figure 7. (a) Thecosomata, Spirellatidae
(Thompson, 1976).
b
(b) Thecosomata, Peraclidae.
(Thompson, 1976).
a
b
Figure 8. (a) Gymnosomata, Crucibranchaea
macrochira (Thompson, 1976).
(b) Gymnosomata, Clione limacina.
(Thompson, 1976).
a
b
Figure 9. (a) Nudibranchia, Polycera capensis
(Thompson, 1976).
8
(b) Nudibranchia, Tritonia festiva
(Thompson, 1976).
In most books and surveys only five orders are treated. Usually Acochlidiacea is not treated
since they are minute and difficult to collect (requires sieving through sand). Thecosomata and
Gymnosomata are also difficult to collect since they are pelagic. Accordingly, most estimates
of opisthobranch species diversity are based on just five groups. One exception to this is
Thompson (1988) who states that there is “about 3000 species” world-wide. (For more
information regarding opisthobranch diversity, see Table 3.)
Table 3. Review of opisthobranch diversity in chosen areas.
Opisthobranchia (A = Estimations, B = Found species)
A Area
Source
3000 Worldwide
Thompson, 1976
700 Tropical Western Marshall & Willan, 1999
Pacific
600 Pacific
Marshall & Willan, 1999
"Equatorial Belt"
550 Great Barrier
Reef
350 S. Barrier Reef Marshall & Willan, 1999
120 Sweden
Hansson, 1998
B Area
Source
561 Madang (PNG) Gosliner (in M & W, 1999)
462 Great Barrier
Reef
439 Guam
Carlson & Hoff (in M & W,
1999)
390 South Africa
Gosliner, 1987
389 Marshall Islands Johnson & Boucher (in M
& W, 1999)
240 Suva (Fiji)
Brodie & Brodie ( in M &
W, 1999)
262 Heron Island
178 Norway
Moen & Svensen, 1999
156 British Isles
Thompson, 1988
105 New Caledonia Risbec, 1928 (in M & W,
1999)
Remarks:
Not including: Acochlidea, Thecosomata, Gymnosomata
Eight orders
Remarks:
Checklist of Great Barrier Reef Opisthobranchia (eight
orders)
Not including Acochlidea
Not including: Thecosomata, Gymnosomata
All opisthobranchs are hermaphrodite and copulation is normally reciprocal (Thompson, 1988;
Hayward & Ryland, 1995). The reproductive organs are present in all adults on the right-hand
side of the body (Hayward & Ryland, 1995). Most species spawn eggs, which hatch as free
swimming, shelled, veliger larvae (Thompson, 1988; Hayward & Ryland, 1995). In Figure 10
there are some examples of typical spawn masses and in Figure 11 there are examples of
veliger larvae.
9
Figure 10. Examples of opisthobranch spawn masses (Behrens, 1991).
10
Figure 11. Examples of opisthobranch veliger larva (Thompson, 1988).
The shelled veliger larva is normally planktotrophic while those species that hatch as miniature
adults are normally lecitotrophic. There is a greater proportion of these species in polar seas
(Willan & Coleman, 1984)
Internally, the opisthobranchs display detorsion and many are secondarily bilaterally
symmetrical. They have one auricle and one nephridium (Ruppert & Barnes, 1994).
Most opisthobranchs are carnivorous, although sacoglossans and aplysiids are herbivorous
(Hayward & Ryland, 1995).
1.4 Nudibranchia
The largest group within the Opisthobranchia is the “Nudibranchia”. Currently, there are
conflicting views whether Nudibranchia is monophyletic (Wägele & Willan, 2000) or
polyphyletic (Tholleson, 1998). Wägele & Willan (2000) goes even further and propose a new
taxon “Nudipleura” as a monophyletic group, which contains Pleurobranchoidea and
Nudibranchia. As with the opisthobranchs, there are different views on nudibranch systematic
and a short review is presented in Table 4. (A historical summary of the nudibranch divisions is
shown in Table 5.)
11
12
Familia
Subfamilia
Tribus
Genus
Subgenus
Species
Superfamilia
Note: Nudibranchia =
Anthobranchia (=
Euctenidieacea =
Ctenidiacea) +
Cladobranchia (=
Actenidiacea)
Guverned by ICZN
Aeolidina
Subordo
Guverned by ICZN
Arminina
Subordo
Doridoidea
Dendronotoidea
Arminoidea
Aeolidoidea
Dendronotina
Subordo
Infraordo
Doridina
Subordo
Marshall & Willan, Rudman & Willan,
1999
1998 (In Beesley et
al., 1998)
* Eudoridoidea
* Polyceroidea
* Anadoridoidea
* Name?
* Name?
* Name?
# Dendronotoidea
# Arminoidea
# Aeolidoidea
Guverned by ICZN
Notaspide(in)a =
Pleurobranchomorph(in)a
Anthobranchi(n)a =
Dorid(ac)ina *
Cladobranchina #
Hansson, 1998
Footnote; ICZN = International Code of Zoological Nomenclature (Fourth Edition 2000)
Note: New taxon
proposed
"Nudipleura" =
Nudibranchia +
Pleurobranchoidea
Anthobranchia
Cladobranchia
Wägele & Willan,
2000 Clades!
Table 4. Systematic review of Nudibranchia according to various authors.
Guverned by ICZN
Aeolidacea
Arminacea
Dendronotacea
Doridacea
Debelius, 1996
Thompson, 1988
Note: In Doridoidea
all species are
holohepatic.
Guverned by ICZN
Aeolidoidea
Arminoidea
Dendronotoidea
Doridoidea
Wells & Bryce,
1993
Note: Doridacea is
divided in three
groups:
cryptobranchs,
porostomes and
phanerobranchs
Guverned by ICZN
Aeolidacea
Arminacea
Dendronotacea
Doridacea
Behrens, 1991
Gosliner, 1987
Note: Sometimes
the Anthobranchia
is divided into
cryptobranchs and
phanerobranchs
Guverned by ICZN
* Doridacea
# Dendronotacea
# Arminacea
# Aeolidacea
Cladobranchia #
Anthobranchia *
Willan & Coleman,
1984
Table 5. Historical summary of the nudibranch divisions (Wägele & Willan, 2000).
The most difficult nudibranch group to characterise is the Arminina/Arminacea/Arminoidea
group (Figure 12).
a
b
Figure 12. (a) Arminina, Armina loveni
(Todd, 1981).
(b) Arminina, Dirona albolineata
(Thompson, 1976).
The other nudibranch groups are somewhat easier to describe as seen in Figures 13-15.
13
a
b
Figure 13. (a) Doridina, Ceratosoma cornigerum
(Thompson, 1976).
a
(b) Doridina, Plocamopherus ceylonicus
(Thompson, 1976).
b
Figure 14. (a) Dendronotina, Doto yongei
(Thompson, 1976).
(b) Dendronotina, Hancockia burni
(Thompson, 1976).
a
b
Figure 15. (a) Aeolidina, Pseudovermis mortoni
(Thompson, 1976).
14
(b) Aeolidina, Facelina annulicornis
(Thompson, 1976).
As with the opisthobranchs, the number of nudibranch species is largest in the tropics (Willan
& Coleman, 1984). As most recent publications treat Nudibranchia as an order or a suborder,
the corresponding lower levels are either suborders or (super) families. Regardless,
Nudibranchia is usually considered to contain four subgroups:
Doridina/Doridacea/Dorodoidea, Aeolidina/Aeolidacea/Aeolidoidea,
Arminina/Arminacea/Arminoidea and Dendronotina/Dendronotacea/Dendronotoidea. The
group with the largest number of species is the Doridina/Doridacea/Dorodoidea. In “Checklist
of Great Barrier Reef Opisthobranchia” (Marshall & Willan, 1999) 281 nudibranch species are
recorded totally and out of them, 204 species belong to the Doridina/Doridacea/Dorodoidea
group. According to Wägele & Willan (2000) Nudibranchia consists of about 190 genera and
60 families. In Table 6, examples of various nudibranch estimations or records are shown.
Table 6. Review of nudibranch diversity in chosen areas.
Nudibranchia (A = estimations, B = found species)
A
3000
800
B
281
Area
Worldwide
Tropical Indo Pacific
Area
Great Barrier Reef
Source
Willan & Coleman 1984
Source
269 Australia
250 Mediterreanean
Cattaneo-Vietti et al, 1990
241 South Africa
Gosliner, 1987
108 British isles
108 British Isles
80 Sweden
Picton & Morrow, 1994
Thompson, 1988
Hansson, 1998
Remarks
Remarks
Checklist of Great Barrier Reef
Opisthobranchia
Checklist of Australian Nudibranchia
Checklist of the Mediterranean
Nudibranchs
Southern African Opisthobranch
Species
British Check List
British Check List
Included 2 species of Notaspidea!
The life span of different nudibranch species varies between a few weeks and two years.
Usually three ecological groupings are used: “Subannual“, “Annual”, and “Biennial” species.
The subannual species are normally feeding on seasonally variable resources while the biennial
species are feeding on stable diets. All nudibranchs appear to be semelparous (Todd, 1981).
The majority of anthobranchs eat sponges or bryozoans and most cladobranchs eat
coelenterates. Other nudibranch diets include ascidians, barnacles, crustaceans, and even the
spawn of other opisthobranchs (Willan & Coleman, 1984). The most complete review
regarding nudibranch food has been provided by McDonald & Nybakken (1997).
Two families of the Doridina/Doridacea/Dorodoidea group are very common both in species
number and abundance in the Indo-Pacific: Chromodorididae and Phyllidiidae.
1.5 Chromodorididae
This family has a lot of species (more than 300 according to Wells & Bryce, 1993) and many
are difficult to distinguish. For instance, in order to separate the genus Chromodoris and the
genus Hypselodoris, the researcher usually has to dissect, prepare and compare the radula
(Figure 16), which is present in all chromodorids. This takes a lot of skill and access to a good
microscope or even a scanning electron microscope (SEM).
15
Figure 16. Examples of opistobranch radular teeth (Photos taken by Terry Gosliner, in Behrens
1991).
Luckily, after this has been done it is often possible to use minor differences in external form
and colour to identify many species. One key characteristic is the shape and form of the
rhinophores (Figure 17).
16
Figure 17. Various shapes and forms of ophistobranch rhinophores (Behrens, 1991).
Especially within the Chromodorididae there are many “colour groups” which are very similar
in appearances (Rudman, 1973, 1977, 1982, 1984, 1991; Gosliner & Johnson 1999). Often
their appearance is quite colourful and it is thought that this is an example of aposematism. As
several species then share the same general colour markings it is suspected that this is also an
example of Müllerian mimicry and that the species take advantage of similar coloration in
order to avoid predation. The deterrent, which makes this work, is normally a substance
acquired from sponges, which are the normal food for chromodorids. In some cases it is just a
question of accumulating useful compounds from the sponges but there are also some
chromodorids which can convert ingested substances to a more “toxic” state (Karuso, 1987).
17
1.6 Phyllidiidae
Another interesting family is the Phyllidiidae, it is also a member of the
Doridina/Doridacea/Dorodoidea group and according to Wells & Bryce (1993) the family
consists of about 70 species. The phyllidiids lack a radula (Brunckhorst, 1993; Yonow, 1996).
Instead they “suck” in pieces of sponges which they utilize as food. Together with the
dendrodorids (which also lacks radula) the phyllidiids constitutes a group which is called
“porostomata”. The phyllidiids can secrete a “milky” fluid when handled and this fluid can
work as a poison. This was first noted when a phyllidiid accidentally killed all the lobsters in
an aquarium where it was put for storage. After this discovery a lot of research have been
conducted on chemical substances in different species of phyllidiids (Karuso, 1987). (Some
chemical structures of these compounds are shown in Figure 18-19).
Figure 18. Toxin from Phyllidia varicosa,
9-isocyanopupukeananae
and its 2-isomer (Karuso, 1987).
Figure 19. Major metabolite from
P. pulitzeri, axisonitrile-1 (Karuso, 1987).
The phyllidiids are mainly restricted to tropical areas and there are just a few species of this
family in more temperate water (Gosliner & Behrens, 1988).
1.7 References
Avila, C. 1995. Natural products of opisthobranch molluscs: a biological review.
Oceanography and Marine Biology: an Annual Review 33: 487-559.
Beesley, P. L., Ross, G. J. B. & Wells, A. (eds). 1998. Mollusca: The Southern Synthesis.
Fauna of Australia. Vol. 5. CSIRO Publishing. Melbourne.
Behrens, D. W. 1991. Pacific Coast nudibranchs: a guide to the opisthobranchs, Alaska to
Baja California. Seachallenger. Monterey.
Bohlin, L. 1989. Pharmacologically active compounds from marine organisms. Acta
Pharmacetica Nordica 1: 175-182.
Brunckhorst, D. J. 1993. The systematics and phylogeny of phyllidiid nudibranchs
(Doridoidea). Australian Museum. Sydney.
Cattaneo-Vietti, R., Chemello, R. and Gianuzzi-Savelli, R. 1990. Atlas of Mediterranean
nudibranchs. La Conchiglia. Roma.
Cimino, G., Fontana, A. and Gavagnin, M. 1999. Marine Opisthobranch Molluscs:
Chemistry and Ecology in Sacoglossans and Dorids. Current Organic Chemistry 3: 327-372.
Debelius, H. 1996. Nudibranchs and sea snails Indo Pacific field guide. IKAN –
Unterwasserarchiv. Frankfurt.
Faulkner, D. J. and Ireland, C. 1977. The chemistry of some opisthobranch molluscs. Pp 2334 in D. J. Faulkner and W. H. Fenical (eds). Marine natural products chemistry. Plenum
Press. New York.
18
Faulkner, D. J. and Ghiselin, M. T. 1983. Chemical defence and evolutionary ecology of
dorid nudibranchs and some other opisthobranch gastropods. Marine Ecology – Progress
Series 13: 295-301.
Gosliner, T. 1987. Nudibranchs of Southern Africa. Seachallenger. Monterey.
Gosliner, T. M. and Behrens, D. W. 1988. A Review of the Generic Divisions Within the
Phyllidiidae with the Description of a New Species of Phyllidiopsis (Nudibranchia:
Phyllidiidae) from the Pacific Coast of North America. The Veliger 30: 305-314.
Gosliner, T. M. and Behrens, D. W. 1990. Special Resemblance, Aposematic Coloration and
Mimicry in Opisthobranch Gastropods. Pp 127-138 in M. Wicksten (ed). Symposium on the
Adaptive Significance of Colour in Invertebrates. University Press Texas A & M.
Gosliner, T. M. and Johnson, R. F. 1999. Phylogeny of Hypselodoris (Nudibranchia:
Chromodorididae) with a review of the monophyletic clade of Indo Pacific species,
including descriptions of twelve new species. Zoological Journal of the Linnean Society
125: 1-114.
Gunthorpe, L. and Cameron, A. M. 1987. Bioactive properties of extracts from Australian
dorid nudibranchs. Marine Biology 94: 39-43.
Hansson, H. G. 1998. Sydskandinaviska marina flercelliga evertebrater, utgåva 2.
Länsstyrelsen Västra Götaland. Göteborg.
Hayward, P. J. & Ryland, J. S. (eds). 1995. Handbook of the Marine Fauna of North – West
Europe. Oxford University Press. Oxford, New York, Tokyo.
ICZN, Fourth Edition. 1999. International Code of Zoological Nomenclature. The
International Trust for Zoological Nomenclature. London.
Karuso, P. 1987. Chemical Ecology of the Nudibranchs. Pp 32-60 in Scheuer, P. J. (ed).
Bioorganic Marine Chemistry, Volume 1. Springer- Verlag. Berlin, Heidelberg.
McDonald, G. & Nybakken, J. 1997. A Worldwide Review of the food of Nudibranch
Mollusks. The Veliger 40: 157-159.
Marshall, J. G. and Willan, R. C. 1999. Nudibranchs of Heron Island, Great Barrier Reef.
Backhuys Publishers. Leiden.
Moen, F. E. & Svensen, E. 1999. Dyreliv i havet. KOM forlag, Kristiansund.
Newman, L. J., Cannon, L. R. G. and Brunckhorst, D. J. 1994. A new flatworm
(Platyhelminthes: Polycladida) which mimics a Phyllidiid Nudibranch (Mollusca,
Nudibranchia). Zoological Journal of the Linnean Society 110: 19-25.
Picton, B. E. & Morrow, C. C. 1994. A Field Guide to the Nudibranchs of the British Isles.
Immel Publishing. London.
Ponder, W. F. and Lindberg, D. R. 1997. Towards a phylogeny of gastropod molluscs: an
analysis using morphological characters. Zoological Journal of the Linnean Society 119: 83265.
Rudman, W. B. 1973. Chromodorid opisthobranch Mollusca from the Indo-West Pacific.
Zoological Journal of the Linnean Society 52: 175-199.
Rudman, W. B. 1977. Chromodorid opisthobranch Mollusca from East Africa and the tropical
West Pacific. Zoological Journal of the Linnean Society 61: 351-397.
Rudman, W. B. 1982. The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West
Pacific: Chromodoris quadricolour, C. lineolata and Hypselodoris nigrolineolata colour
groups. Zoological Journal of the Linnean Society 76: 183-241.
Rudman, W. B. 1984. The Chromodorididae (Opisthobranchia: Mollusca) of the Indo –West
Pacific: a review of the genera. Zoological Journal of the Linnean Society 81: 115-273.
Rudman, W. B. 1991. Purpose in pattern: the evolution of colour in chromodorid nudibranchs.
Journal of Molluscan Studies 57: 5-21.
Rudman, W. B. 2000. The Sea Slug Forum, www.austmus.gov.au/science/invert/mal/forum
Ruppert, E. E. & Barnes, R. D. 1994. Invertebrate Zoology. Saunders College Publishing.
Fort Worth, Philadephia, San Diego.
19
Thollesson, M. 1998. Nudibranch systematics and molecular data. PhD Thesis, Göteborgs
Universitet.
Thompson, T. E. 1976. Biology of opisthobranch molluscs. Volume I. The Ray Society.
London.
Thompson, T. E. 1988. Molluscs: Benthic Opisthobranchs. E. J. Brill / Dr W. Backhuys.
Leiden, New York, København, Köln.
Todd, C. D. 1981. The ecology of nudibranch molluscs. Oceanography and Marine Biology:
an Annual Review 19: 141-234.
Tullrot, A. 1998. Evolution of warning coloration in the nudibranch Polycera quadrilineata.
PhD Thesis, Göteborgs Universitet.
Wells, F. E. & Bryce, C. W. 1993. Sea slugs of Western Australia. Western Australian
Museum. Perth.
Willan, R. C. & Coleman, N. 1984. Nudibranchs of Australasia. Australasian Marine
Photographic Index. Sydney.
Wägele, H. & Willan, R. C. 2000. Phylogeny of the Nudibranchia. Zoological Journal of the
Linnean Society 130: 83-181.
Yonow, N. 1996. Systematic revision of the family Phyllidiidae in the Indian Ocean province:
Part 1 (Opisthobranchia: Nudibranchia: Dorodoidea). Journal of Conchology 35: 483-516.
20
2 Checklist of opisthobranchs of the Wakatobi National Park, SE
Sulawesi, Indonesia
2.1 Introduction
Operation Wallacea (“Opwall”) is a joint venture between the Indonesian “Wallacea
Development Institute” and a British company: Ecological Surveys Ltd. The co-operation
started 1995 and the purpose was to promote conservation work in a remote Indonesian
archipelago, Tukang Besi (Chart 1). The Operation Wallacea works with both land and marine
projects, concentrating mainly on field surveys regarding diversity issues but it has also been
involved in some projects promoting sustainable use of natural resources. This means, for
instance, changing the fishing methods and promoting ecotourism in the area. A significant
result of this work was achieved when “The Wakatobi Marine National Park” was created in
July 1996. It is currently the second largest Indonesian “National Marine Park” (13000 km2)
stretching between 05º15’S, 123º23’E and 06º08’S, 124º37’E (Stanzel & Newman, 1997). The
name Wakatobi comes from the first letters of the four largest islands in the area (Wanci,
Kaledupa, Tomea and Binongko).
One aim of Operation Wallacea is to survey and publish checklists of the marine life in the
national park. During 1997-99 a lot of work was geared towards a checklist of
Opisthobranchia. This paper presents results from this work.
Chart 1. Tukang Besi archipelago, Indonesia (Expedia, 2001).
2.2 Methods
The marine projects, run by Operation Wallacea, are concentrated around a small island, Hoga
(Chart 2), situated near Kaledupa. During my visit 1999, two major projects were conducted,
one regarding correlation between butterfly fish diversity and coral diversity and another one
regarding opisthobranch diversity. Paying volunteers did nearly all work but there was also
some scientific staff supervising the projects. The opisthobranch project started in 1997 (two
21
weeks) and continued for 10 months in 1998 and for 7 months in 1999. The main effort was
done during 1999 when a lot of work was concentrated at opisthobranchs.
Hoga Island
Chart 2. Hoga Island, Tukang Besi archipelago, Indonesia (Expedia, 2001).
In the opisthobranch project, groups of diving volunteers concentrated on finding as many
species of opisthobranchs as possible (down to a maximum depth of 30 m). For each recording
they noted numbers of individuals, dates, depths and sizes. All species were photographed and
unidentifiable species were also drawn by hand. The main identification books used at Hoga
were: Allen & Steene, 1996; Behrens, 1991; Coleman, 1989; Debelius, 1996; Gosliner, 1987,
Gosliner, et al, 1996; and Wells & Bryce, 1993. The collected data for 1999 (handwritten
notes) were then taken to England and later also copied and sent to Sweden where the
compilation (Appendix I) was made during the year of 2000. During this work it was possible
to identify some more species with the help of the Internet, especially as the site “The Sea Slug
Forum” proved to be very valuable (Rudman, 2000).
At least one specimen of each species was conserved at Hoga for later shipment to the Bogor
Zoological Museum. Also, from many species, a small tissue sample (large species) or a whole
individual (small species) was taken and conserved. The aim with this material is to compare
the DNA from the species and to do a cladistic analysis later on (Warren, 2000 and personal
communication, October 1999*).
Notes of the taxa found during 1997 or 1998 but not 1999 are added to “Appendix I” but these
taxa are not included in calculations regarding “observations” or “occasions” due to lack of
data. The reason for choosing the word “observations” instead of “individuals” is because each
individual could have been recorded more than once. The term “occasions” is used since not all
survey work was done diving. In a few cases surveys were made in the intertidal area during
low tide.
* Lindsay Warren, personal communication, October 1999 (Hoga Island).
22
2.3 Results
A total of 60 different species were found during two weeks in September 1997. Between
March and December 1998 another 115 opisthobranch species were found. The total species
number for the whole period 1997-1999 is 297 species (Warren, 2000).
During the season of 1999 (May-November), 3523 opisthobranchs were found during the
survey. They belonged to a total of 235 identified or suspected (provisionally identified)
species and to 73 identified genera (Appendix I).
Table 1. Summary of “The 1999 Opisthobranch Survey – Operation Wallacea” (Indonesia)
Anaspidea Cephalaspidea Notaspidea Nudibranchia Sacoglossa
Species
9
36
5
161
23
Observations 116
316
42
2888
159
Two additional observations were of Colpodaspis thompsoni, which probably belong to
Cephalaspidea but its current taxonomic status is uncertain (Marshall & Willan, 1999). The
results regarding discovered species and genera for the periods 1997-1999, 1997-1998 and
1999 are shown in Figure 1 and 2.
Operation Wallacea
Sacoglossa
Category
Nudibranchia
1997-1999
1999
1997-1998
Notaspidea
Cephalaspidea
Anaspidea
0
50
100
150
200
250
Number of species
Figure 1. Summary of species, “Operation Wallacea - Opisthobranchia survey” 1997-1999
(Indonesia).
23
Operation Wallacea
Sacoglossa
Category
Nudibranchia
1997-1999
Notaspidea
1999
1997-1998
Cephalaspidea
Anaspidea
0
20
40
60
Number of identified genera
Figure 2. Summary of identified genera, “Operation Wallacea - Opisthobranchia survey”
1997-1999 (Indonesia).
In Figure 3 there is a summary of the results of the identification effort.
Identification of Opisthobranchs 1997-99
Genus + species
Genus
Only higher taxa
Figure 3. Result of identification effort regarding 295 opisthobranch taxa, “Operation
Wallacea - Opisthobranchia survey” 1997-1999 (Indonesia).
The proportions (species and genera) between the nudibranch suborders for the period 19971999 are presented in Figure 4 and 5.
24
Nudibranchia 1997-99
Aeolidina
Arminina
Dendronotina
Doridina
Figure 4. Proportion of species within nudibranch suborders, Operation Wallacea survey 19971999 (Indonesia). Totally 208 nudibranch species.
Nudibranch genera 1997-99
Aeolidina
Arminina
Dendronotina
Doridina
Figure 5. Proportion of genera within nudibranch suborders, Operation Wallacea survey 19971999 (Indonesia). Totally 56 nudibranch genera.
25
Table 2 presents a list of the most common species. Some of these species (and a few others)
are illustrated in Plate A and B.
Table 2. The most common opisthobranch species during Operation Wallacea survey 1999
(Indonesia).
Order
Species
Observations Occasions
Cephalaspidea
Notaspidea
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Nudibranchia
Sacoglossa
Sacoglossa
Sacoglossa
Chelidonura amoena
Pleurobranchus forskalii
Phidiana indica
Phyllodesmium briareum
Pteraeolidia ianthina
Dermatobranchus sp (OWN 101)
Chromodoris annae
Chromodoris coi
Chromodoris dianae
Chromodoris geometrica
Chromodoris kuniei
Chromodoris leopardus
Chromodoris lochi
Chromodoris magnifica
Chromodoris willani
Hypselodoris bullocki
Jorunna funebris
Nembrotha kubaryana
Phyllidia coelestis
Phyllidia elegans
Phyllidia ocellata
Phyllidia varicosa
Phyllidiella pustulosa
Phyllidiella rudmani
Phyllidiopsis pipeki
Phyllidiopsis shireenae
Phyllidiopsis striata
Reticulidia fungia
Roboastra gracilis
Elysia sp (OWN 99)
Thuridilla bayeri
Thuridilla hoffae
Total number
24
20
17
398
25
20
215
35
106
49
22
13
132
119
106
18
29
36
113
164
19
131
604
30
58
12
34
15
16
15
36
13
2644
17
11
17
27
19
13
109
30
60
40
17
11
81
71
63
13
26
?
82
115
19
96
289
24
52
11
28
14
16
12
25
11
2.4 Discussion
Appendix I is the summary of more than 450 hand-written pages with opisthobranch
information from the 1999 survey. The material was delivered in three batches between
January and April 2000. As numerous people wrote the material, a continuing obstacle was to
decipher the notes accurately. Another difficulty was the uses of many synonyms as this made
the compilation work very time consuming. A third problem was the fact that the most
complete photographic handbook (Debelius, 1996) contains many misidentifications and lacks
information about authors.
26
As seen from Figure 1 most of the species (in all orders) were found during 1999. The reason
for this is probably because of the increased effort under the 1999 season (May – November).
Regrettably, there is no information regarding search effort, which makes it difficult to
compare the three years.
When looking at the number of genera instead (Figure 2) the picture is less clear. Fewer
Cephalaspidean genera were found 1999 than 1997-1998. This is probably because of less
search effort during 1999 than 1997-1998. (During my own visit, no actual digging was
conducted in search of the burrowing Cephalaspidea.)
It is interesting to note that, despite a massive identifying effort, only about 60 % of the
opisthobranchs were identified to species level (Figure 3).
The majority of the observations involved members of the Nudibranchia, which is not
surprising since Nudibranchia is the largest order in Opisthobranchia. As seen from Figures 4
and 5, the most common nudibranch suborder is the Doridina whether looking for species
number or generic number.
There are very few similar surveys to compare with. The most recent one is the “Nudibranchs
of Heron Island, Great Barrier Reef” (Marshall & Willan, 1999). This survey was run between
November 1980 and December 1998 and a total of 262 species was found during that period.
In comparison 295 species were found during the Operation Wallacea surveys 1997-1999
(Figures 1 and 3). This number will probably be somewhat lower after final examinations but it
is still an impressive figure after only three years of surveys.( According to Warren (2000)
there are 297 species but I have excluded Colpodaspis thompsoni and Tritonopsilla australis.
In the first case because of its uncertain taxonomic status and in the later case because I believe
it to be a case of mistaken identity.)
The highest count of opisthobranchs for an ongoing survey in the Indo Pacific is 561 species
(Madang, Papua New Guinea) but that is for the period 1986 – March 1997 (Gosliner in
Marshall & Willan, 1999). If not just looking at surveys then Gosliner & Draheim (1996) give
the highest count of Indo Pacific opisthobranchs in a study. They state that there are more than
3400 opisthobranch species in the region and that the locality with the most species, 646, is
Papua New Guinea. No doubt, these figures shows that the region of Papua New Guinea and
Indonesia is the leading diversity centre for Opisthobranchia.
According to Table 2, 75 % of the opisthobranch observations were of the 14 % most common
species. One problem is to categorise the found species. Is a species always common when
found in large numbers? What if the species is found only a few times but in large number? In
my case I choose to regard all species, which were found on more than 10 separate occasions
(during the 1999 season) as common.
In “Nudibranchs of Heron Island, Great Barrier Reef” (Marshall & Willan, 1999) there is an
attempt to give a relative abundance scale. This could function but since the authors do not
give information about the time duration of sampling periods it was not possible to use their
abundance scale in this report.
27
2.5 References
Allen, G. R. & Steene, R. 1996. Indo-Pacific coral reef field guide. Tropical Reef Research.
Singapore.
Behrens, D. W. 1991. Pacific Coast nudibranchs: a guide to the opisthobranchs, Alaska to
Baja California. Seachallenger. Monterey.
Coleman, N. 1989. Nudibranchs of the South Pacific. Neville Coleman’s Sea Australia
Resource Centre. Springwood.
Debelius, H. 1996. Nudibranchs and sea snails Indo Pacific field guide. IKAN –
Unterwasserarchiv. Frankfurt.
Expedia, 2001. Internet site: www.expedia.com (March 2001).
Gosliner, T. 1987. Nudibranchs of Southern Africa. Seachallenger. Monterey.
Gosliner, T. M., Behrens, D. W. and Williams, G. C. 1996. Coral Reef Animals of the IndoPacific. Seachallengers. Monterey.
Gosliner, T. M. and Draheim, R. 1996. Indo-Pacific opisthobranch gastropod biogeography:
how do we know what we don’t know? American Malacological Bullentin 12: 37-43.
Marshall, J. G. & Willan, R. C. 1999. Nudibranchs of Heron Island, Great Barrier Reef.
Rudman, B. 2000. The Sea Slug Forum. (www.austmus.gov.au/science/invert/mal/forum)
Stanzel, K. B. & Newman, H. 1997. Progress report on the 1996 marine survey of the Tukang
Besi (Wakatobi) archipelago, South East Sulawesi, Indonesia. (Internal report: Operation
Wallacea).
Warren, L. 2000. Opisthobranchs of the Tukang Besi archipelago, S.E. Sulawesi, 1999 survey
report & results. (www.operationwallacea.win-uk.net/opisthob/opisthob.htm (2000-11-15))
Wells, F. E. & Bryce, C. W. 1993. Sea slugs of Western Australia. Western Australian
Museum. Perth.
28
3 Spatial distribution of nudibranchs according to benthic structure
3.1 Introduction
Why do we know so little about nudibranch ecology? One reason is that nudibranch
populations are both scarce and transient (Todd, 1981, Marshall & Willan, 1999). This means
that finding places where one can do nudibranch research are difficult. One potential site for
such research is at Hoga Island (Indonesia) as the nudibranchs are rather abundant at this area.
There are also many nudibranch species around Hoga that are quite common and not so
shortlived and that makes it possible to do interesting scientific research at this locality. The
research facility is run by Operation Wallacea, a joint venture between The Wallacea
Development Institute (Indonesia) and Ecosurveys Ltd (United Kingdom).
Hoga Island is situated in the Tukang Besi archipelago, South East of Sulawesi. In July 1996
the area was designated as the Wakatobi Marine National Park (Stanzel & Newman, 1997) and
since then there has been ongoing research at this locality.
My hypothesis was:
Nudibranch distribution depends on benthic structure.
The reason for the hypothesis is that I wanted to see if it was possible to find environment
preferences among some of the more common nudibranchs in the area.
3.2 Material and methods
The dive sites, chosen for research, were “Sampela” and “Buoy 3” (Chart 1). Sampela was
considered an example of a more polluted area (from a nearby village) and Buoy 3 an example
of a more “unspoilt” area. The survey was conducted in 1999 between the eight and twelfth of
November (five consecutive days).
On both locations we put down two stationary lines, that each were 30 m long. One line was
put at “+12 m” and another one at “+3 m”. The “+” sign indicates that the lines were put down
on an estimated depth of 3 or 12 m below the reef crest. This was done in accordance with
other ongoing projects and relevant literature at hand (English et al. 1994). The actual survey
depth varied with the tidal state but we recorded the depth of the starting point on each dive so
that we could adjust our measurements later if necessary. On each dive we aimed to spend 20
minutes searching along each line. The search area was roughly 2.5 m above and under the
line, thus having a search area of about 150 m2. An alternative search description is to say that
we surveyed a volume of 300 m3 (this figure is calculated from the volume of a half cylinder
with a radius of 2.5 m). We dived at each line twice a day (before and after lunch), recording
observed species, depths and numbers and measuring individual sizes. The number of divers,
on each dive, varied between two and five.
29
Buoy 3
Sampela
Chart 1. Location of the dive sites Buoy 3 and Sampela (Tukang Besi, Indonesia)
In order to measure the benthic structure, the length of the benthic forms along the line transect
was recorded. Standardised codes were used to describe the benthic forms, see Table 1.
Collected data were later analysed by using statistical methods such as ANOVA (Analysis of
Variance) and DCA (Detrended Correspondence Analysis). Comparable indices regarding
benthic structure and diversity of nudibranchs for the two sites were computed by using the
Shannon-Weaver Index formula: H = - Σ pi ln pi (Fowler et al., 1998). (The term pi is the
proportion of a particular species in a sample which is multiplied by the natural logarithm of
itself. H is derived by summing the product for all species in the sample. The minus sign is to
make the final value of H positive.)
30
Table 1. Standardised benthic codes. After English et al., 1994.
ACB
ACE
ACD
ACS
ACT
CB
CD
CE
CF
CM
CMR
CS
DC
DCA
CHL
CME
=
=
=
=
=
=
=
=
=
=
=
=
=
=
=
=
Hard coral
Acropora
Acropora
Acropora
Acropora
Acropora
Coral
Coral
Coral
Coral
Coral
Coral
Coral
Dead coral
Dead coral
Heliopora
Millepora
Branching
Encrusting
Digitate
Submassive
Tabulate
Branching
Digitate
Encrusting
Foliose
Massive
Mushroom
Submassive
AA
CA
HA
MA
TA
=
=
=
=
=
DDD
R
RCK
S
SI
WA
=
=
=
=
=
=
Algea
Algal assembly
Coralline algea
Halimeda
Macro algea
Turf algea
Abiotic
Data missing
Rubble
Rock
Sand
Silt
Water
Other
Other
Soft coral
Sponge
Sponge
Sponge
Sponge
Sponge
Sponge
Sponge
Sponge
Sponge
OT
SC
SP
SPB
SPD
SPE
SPF
SPM
SPO
SPS
SPT
=
=
=
=
=
=
=
=
=
=
=
ZO
= Zoanthids
Branching
Digitate
Encrusting
Foliose
Massive
Other form
Submassive
Tubular, Barrel,
Vase
With algea
3.3 Results
The results from measuring benthic structure are shown in Table 2. Shannon-Weaver Indices
for the four transects were calculated according to Fowler et al., 1998.
Table 2. Benthic structure at Buoy 3 and Sampela (Tukang Besi, Indonesia).
Benthic code
Buoy 3
Buoy 3
Sampela
Sampela
(See Table 1)
(“+ 3 m”) (“+ 12 m”) (“+ 3 m”) (“+ 12 m”)
% Cover
% Cover
% Cover
% Cover
AA
1
2
1
3
ACB
3
CA
7
32
9
20
CB
10
1
CD
10
1
CE
13
15
7
7
CF
2
1
1
CM
1
CMR
3
4
CS
10
DCA
3
3
DC
7
MA
1
OT
2
1
R
12
2
25
11
S
3
38
SC
3
7
13
12
SP
2
5
7
3
SPE
1
4
WA
26
32
3
2
Shannon2.251
1.746
2.422
1.780
Weaver Index
31
The average line depth and search time (and their standard deviation = SD) for Buoy 3 and
Sampela are given in Table 3.
Table 3. Average search time and line depth for Buoy 3 and Sampela (Tukang Besi,
Indonesia).
Buoy 3 Buoy 3 Buoy 3 Buoy 3
Sampela Sampela Sampela Sampela
AM
PM
AM
PM
AM
PM
AM
PM
“+3m” “+3m” “+12m” “+12m” “+3m” “+3m” “+12m” “+12m)
Average
27
27
30
31
32
28
33
31
search
time
(min)
SD
8
3
11
14
7
6
13
14
Average
4.7
4.7
11.9
14.0
5.5
6.8
12.2
13.3
line
depth
(m)
SD
0.78
0.18
0.22
0.48
0.24
0.05
0.41
0.25
During 40 dives we managed to find a total of 252 nudibranchs belonging to at least 19 species
(two nudibranch taxa were not identified to species level). At Sampela we found 138
nudibranchs and at Buoy 3 we found 114. The search record regarding observations and
species is presented in Table 4. The complete search record is available in Appendix II.
Table 4. Search record for Buoy 3 and Sampela (Tukang Besi, Indonesia).
Species
Buoy 3
Sampela
15
7
Chromodoris annae
3
0
Chromodoris boucheti
0
5
Chromodoris coi
6
0
Chromodoris dianae
5
0
Chromodoris elisabethina
1
2
0
2
Chromodoris kunei
8
0
19
0
17
21
Jorunna funebris
6
0
Nembrotha kubaryana
3
0
Reticulidia fungia
5
7
0
9
3
15
Phyllidia coelestis
0
2
Phyllidia sp
5
10
Phyllidia varicosa
18
55
Phyllidiella pustolosa
Unidentified
0
1
0
1
Pteraolidia ianthina
0
1
Thecacera picta
The Shannon-Weaver Index (Fowler et al., 1998) regarding nudibranch species for Buoy 3 is
2.377 and for Sampela 1.976.
32
The effects of the sites (Sampela, Buoy 3), depths (+3m, +12m) and times (“am”, “pm”) were
analysed by an ANOVA (By the computer program MANOVA). Both nudibranch
observations (Figure 1) and species number (Figure 2) were tested. The lowest p-value (0.08)
was found for interaction of sites and depths regarding number of species. In no cases “am/pm”
seemed to affect species number or observations.
B u o y 3 - S a m p e la
90
Number of observations
80
70
60
T o ta l B u o y 3
50
A ve ra g e B o u y 3
T o t a l S a m p e la
40
A v e r a g e S a m p e la
30
20
10
(+12m)
Phyllidiidae
(+12m)
Chromodorididae
Totally (+12m)
(+3m)
Phyllidiidae
(+3m)
Chromodorididae
Totally (+3m)
0
Figure 1. Buoy 3 and Sampela (Tukang Besi, Indonesia): Nudibranch observations and
average (with SE bars) at two depths.
16
Number of species
14
12
B uo y 3
A ve ra g e B u o y 3
10
8
S a m p e la
A ve ra g e S a m p e la
6
4
2
(+12m)
Phyllidiidae
(+12m)
Chromodorididae
Totally (+12m)
(+3m)
Phyllidiidae
(+3m)
Chromodorididae
Totally (+3m)
0
Figure 2. Buoy 3 and Sampela (Tukang Besi, Indonesia): Species number and average (with
SE bars) of nudibranchs, at two depths.
33
14
Species number
12
10
8
T o ta l
6
A ve ra g e
4
(Sampela)
Phyllidiidae
(Sampela)
Chromodorididae
Totally (Sampela)
(Buoy 3)
Phyllidiidae
(Buoy 3)
Chromodorididae
0
Totally (Buoy 3)
2
Figure 3. Buoy 3 and Sampela (Tukang Besi, Indonesia): Total species number and average
(with SE bars) of nudibranchs.
Collected data was also used in a Detrended Correspondence Analysis (DCA). This is an
indirect gradient analysis technique which can be used for explanatory analysis (Gauch, 1984).
The analysis was made with the computer program CANOCO 4.0 (Ter Braak, 1998). The
result from this analysis is shown in Figure 4.
34
4
Chromodoris dianae
Chromodoris annae
2
Phyllidia sp
Chromodoris coi
Phyllodiopsis striata
Chromodoris kunei
Jorunna funebris
Reticulidia fungiaChromodoris boucheti
Phyllidia coelestis
Chromodoris elisabethinaThecacera picta
Phyllidia varicosa
0Nembrotha kubyaryana
Ptereoalidia ianthina
Sp
0
2
4
6
1.8
Macro algae
Cor. digitate
Cor. branching
1.6
Other
Cor. foliose
Dead cor. with algae
Bouy 3
Water
1.4
+3
[Cor. encr.,Cor. mushroom ]
PM
AM Rubble
Algal assembly
Cor. algea Soft coral
Sponge +12
Sand
Sponge encrusting
Sampela
[Acropora encr,Cor. submassive,Dead coral ]
1.2
Cor. massive
1
2
3
Figure 4. Results from a “Detrended Correspondence Analysis” (DCA). The data are taken
from the sites: Buoy 3 and Sampela (Tukang Besi, Indonesia)
3.4 Discussion
Totally, the average number of observed nudibranchs was higher at Sampela (7) than Buoy 3
(6). We found approximately the same average number (6) of nudibranchs around Buoy 3
regardless of depth. At Sampela there was a difference in nudibranch abundance between the
35
depths. We found on average more nudibranchs at “+3m” then at “+12m” (Figure 1). The
lower number for “+12m” is probably due to the fact that there was a large sandy area at a part
of this depth (very few nudibranchs live on sand).
At both depths at Buoy 3 the normal pattern was that we would find more chromodorids than
phyllidiids. This was not the case at Sampela however, where the situation was reversed. As
this reversal occurs whether we look at individual number (Figure 1) or number of species
(Figure 2), it would be interesting to check more similar sites and see if there is a larger pattern.
There is a possibility that this reversal could work as an environment indicator since we
consider Sampela more polluted than Buoy 3.
The highest number of species, 14, was found at Buoy 3 but Sampela was not far behind with
12 species. This was in total numbers, when looking at the average values per dive there was
no difference between the sites. On each dive we could expect to find about 4 species whether
we were diving at Buoy 3 or Sampela (Figure 3).
When comparing the “Shannon-Weaver Index” for the two sites (Sampela, 1.976 and Buoy 3,
2.377) there is a cause to say that Buoy 3 seemed to be more diverse than Sampela.
As for my hypothesis that “Nudibranch distribution depends on benthic structure”, it is difficult
to get a clear picture of what kind of benthic structure the different nudibranchs prefer but
according to Figure 4 some of the surveyed nudibranch species are affiliated with specific
benthic forms, as Chromodoris dianae, C. annae and C. magnifica occurred with macro algea,
digitate and branching coral. Another indication is that Chromodoris dianae, C. annae, C.
magnifica, C. coi, C. kunei, Phyllidiella pustolosa, Phyllidia sp and Phyllidiopsis striata
seemed to prefer “+3m” to “+ 12m”.
Since most nudibranchs are supposed to settle near food sources, I had expected a much more
clear and close affiliation between Chromodoridae/Phyllidiidae and the “sponge” categories, as
sponges are supposed to be their food source. This connection is not so easy to see in Figure 4.
Nevertheless, I still think it should be possible to find out more facts regarding nudibranchs
and benthic structure, by doing more survey attempts like the one presented here.
3.5 References
English, S., Wilkinson, C. and Baker, V. (eds). 1994. Survey Manual for Tropical Resources.
Australian Institute of Marine Science. Townsville.
Fowler, J., Cohen, L. and Jarvis, P. 1998. Practical Statistics for Field Biology. John Wiley
& Sons. Chichester, New York, Weinheim, Brisbane, Singapore, Toronto.
Gauch, H. G. Jr. 1984. Multivariate analysis in community ecology. Cambridge University
Press. Cambridge, London, New York, New Rochelle, Melbourne, Sidney.
Marshall, J. G. & Willan, R. C. 1999. Nudibranchs of Heron Island, Great Barrier Reef.
Stanzel, K. B. & Newman, H. 1997. Progress report on the 1996 Marine Survey of the
Tukang Besi (Wakatobi) archipelago, South East Sulawesi, Indonesia. (Internal report:
Operation Wallacea).
Ter Braak, C. J. F. 1998. Canoco 4.0 (Computer program). Centre for Biometry Wageningen.
Wageningen.
Todd, C. D. 1981. The Ecology of Nudibranch Molluscs. Oceanography and Marine Biology:
an Annual Review 19: 141-234.
36
4 Conclusions and recommendations
In section 3, I had another hypothesis: Chromodorids have larger home ranges than Phyllidids.
The background for this hypothesis was that there was a general field observation among the
volunteers that nudibranchs of the family Phyllidiidae were much more stationary than
nudibranchs of the family Chromodorididae. Accordingly, I wanted to objectively test this
observation.
We first tried to use a coordinate system and write down the co-ordinates for every nudibranch
we found, hoping to be able to follow their movement. It seemed to work at first but later a
storm ruined our efforts by moving the lines a little bit and thus made it impossible to continue
using co-ordinates. We then tried to mark the nudibranch position with a string of rope and a
buoyant bamboo marker. This also seemed to work at first but we later got problem as the
bamboo markers lost their buoyancy and that made it very difficult to find them. It was also
difficult to recognise the specific nudibranchs and I believe that we would have needed an
identification system in order to successfully follow the nudibranchs. Perhaps a “fish marking”
system with tags would work? Another problem is to keep a functioning “dive relay race”
going on for a long enough period.
When doing my own survey we also discovered how difficult it is to do scientific work
underwater. As we had to be several divers sharing the workload we always had to adjust for
the least experienced and/or the diver with the highest air consumption. This makes underwater
surveys with volunteers extremely difficult due to the differences in diving experience,
biological knowledge and air consumption. It takes many dives before you can find a system
that is workable.
Another problem with working in a remote area like Wakatobi is the lack of equipment and the
logistic transport problems when needing repairs or spare parts. This is especially hard since
the climate affects and wears down all kinds of equipment, for instance, diving equipment,
boats, recording devices and of course computer equipment. There is always the need to come
up with new repair ideas, using very simple tools or equipment. We also found out how hard
the tropical environment affects the personnel, usually about one third are on the sick list due
to infections (stomach illness, ear infections, badly infected wounds, etc). This means that you
always have to have a lot more people helping you than you would normally expect.
In the future I think it would be worthwhile to do a more structured opisthobranch study at
Hoga. One main mistake, when working with the checklist (section 2), is that there has been no
registration of search effort. This makes it impossible to correlate the opisthobranch abundance
with, for instance, dive time or search area. The easiest way to register the search effort is to
keep track of the “active search time” but another idea could be to cooperate with other
projects where the search area is defined by transects. In these cases an estimation of search
area is given for “free”!
37
5 Acknowledgements
I would like thank:
Per Milberg, for all the help with this report. - I could not have done it without your help.
Lindsay Warren and Krzystof Brzkieta, for trying to help out at Hoga when my appointed
(Opwall) supervisor disappeared!
All the volunteers at Hoga who graciously tried to help me, especially my dive buddies: Peter
Bächi, Rob Saunders, Nikki Rowland, Jenny Hill, Elaine Worley, Gail Young, Sarah Yeates
and Tony Gatcombe.
Richard Willan, Pam Beesley, The Ray Society, David Behrens and the Linnean Society of
London for kind permissions to reproduce appropriate illustrations in my thesis.
My parents, for financial and moral support.
Åsa Nilsson, my fiancé, for helping me, all the time. – Thank You!
38
Plate A
Chelidonura amoena Bergh, 1905
Pleurobranchus grandis Pease, 1868
Note: White squid cleaning the opisthobranch!
Cyerce elegans Bergh, 1870
Nembrotha sp (lineolata? Bergh, 1905)
Jorunna funebris (Keelart, 1858)
Hexabranchus sanguineus (Rüppell & Leuckart,
1828)
Note: The identification details are taken from “The Sea Slug Forum” (www.seaslugforum.net 2001-0422). Copyright holder and photographer is the author (Lars Karlsson). The photos are all from Hoga Island
(1999).
Plate B
Chromodoris annae Bergh, 1877
Chromodoris magnifica (Quoy & Gaimard, 1832)
Chromodoris sp (boucheti? Rudman, 1982)
Phyllidiella pustulosa (Cuvier, 1804)
Phyllidia sp (willani? Brunckhorst, 1993)
Phyllidiopsis striata Bergh, 1888
Note: The identification details are taken from “The Sea Slug Forum” (www.seaslugforum.net 2001-0422). Copyright holder and photographer is the author (Lars Karlsson). The photos are all from Hoga Island
(1999).
Haminoeidae
Atys
Atys
Atys
Haminoeidae
Atys
Haminoeidae
Haminoeidae
Atys
Atys
Haminoeidae
Haminoeidae
Atys
Atys
Bulla
Bullidae
Bullidae
Bulla
Bulla
Bulla
Bullidae
Chelidonura
Aglajidae
Aglajidae
Chelidonura
Aglajidae
Chelidonura
Aglajidae
Aglajidae
Chelidonura
Aglajidae
Aglajidae
Chelidonura
Chelidonura
Aglajidae
Chelidonura
Chelidonura
Chelidonura
Chelidonura
Chelidonura
Chelidonura
Aglajidae
Chelidonura
Aglajidae
Chelidonura
Aglajidae
Aglajidae
Chelidonura
Aglajidae
Gastropteron
Haminoea
Haminoea
Haminoea
Haminoea
Haminoea
Haminoea
Aglajidae
Haminoea
Aglajidae
Ilbia
Ilbidae
Odontoglaja
Aglajidae
Aglajidae
Smaragdinellidae Smaragdinellidae Phaneropthalmus
Philinopsis
Aglajidae
Aglajidae
Philinopsis
Aglajidae
Aglajidae
Philinopsis
Philinopsis
Sagaminopteron
Gastropteridae
Gastropteridae
Order = Anaspidea
Haminoeidae
cylindrica / cylindricus
cf. cylindricus
debilis
naucum
cf naucum
semistriata / semistriatus
cf semistriatus
sp
ampulla
sp
sp
vernicosa
amoena
cf. amoena
hirundinina
inornata
lineolata
sandrana
sp
sp
sp
sp
sp. 1
sp. 2
sp. 3.
tsurugensis
varians
sp.
sp
sp
sp
sp
sp 5
sp. 1.
sp. 2.
mariana
guamensis
smaragdinus
gardineri
pilsbryi
sp.
sp.
psychedelicum
Aplysiidae
Aplysiidae
Aplysiidae
Aplysiidae
Aplysiidae
Aplysiidae
Aplysiidae
Allan 1932
extraordinaria
sp
sp
sp
auricularia
sp. 1.
longicauda
striata
Aplysia
Aplysia
Aplysia
Aplysia
Dolabella
Phyllaplysia
Stylocheilus
Stylocheilus
Aplysiid
Aplysiidae
Aplysiidae
3
1
1
4
1
1
1
5
1
3
1
1
1
2
8
4
1
2
1
1
8
40
1
1
21
1
1
1
5
1
4
1
1
1
2
14
85
1
2
1
1
9
Rudman, 1973
Carlsson & Hoff, 1974
x
x
Hoff & Carlsson, 1990
Rudman, 1978
(Ruppell & Leuckart, 1828)
(Eliot, 1903)
(Eliot, 1900)
x
x
Baba & Abe, 1959
Eliot, 1903
17
1
5
5
1
4
3
4
1
1
2
1
2
2
2
2
1
8
8
1
6
3
24
1
7
14
1
8
10
37
1
1
11
1
2
116
2
3
2
1
21
39
3
42
3
970920
981124
981124
981119
981001
980907
970920
980620
980621
981124
970920
981028
980501
980519
980519
980519
980901
970920
980901
980524
980530
980901
980801
Obser- Occa- First
vations sions 97/98
Gould, 1859
Bergh, 1905
x
(Quoy & Gaimard, 1832)
Baba, 1949
Linnaeus, 1758
Pease, 1860
Pease, 1860
(Linnaeus, 1758)
(Helbling, 1779)
(Lightfoot, 1786)
x
(Qouy & Gaimard, 1825)
Authors
Species
(If identified)
Genus
(If identified)
Family
"CGBRO" **
Family
"SSF" *
Appendix I, 1(6)
Compilation over Opisthobranchs found during Operation Wallacea survey 1997-1999.
990928
990911
990730
991116
990825
9906
990925
990825
990621
990931
990917
991031
990929
990611 991114
990528
990728 990715
990524
990617
990604 991101
990615
990615
990511
990730
991001
990725
990614
991103
990524
990519
990621
990729
990616
990617
990725 990929
990515 991101
991027
990702 990929
990917 991105
990708
990906 991102
991103 991116
991102 991116
990730
95
116
52
204
98
97
122
152
115
78
79
70
26
28
29
113
114
89
73
209
38
200
202
166
32
33
141
182
178
164
110
184
156
172
121
Sombano lake (83).In "GBRO" (Rüppell & Leuckart, 1830)
See SSF under Haminoea sp. 3.
See SSF under Haminoea sp. 3.
OS0775?
Probably Philinopsis lineolata
In "GBRO": (Quoy & Gaimard, 1833)
In "GBRO": Linné, 1758
semistriatus
(semistriata?)
In "GBRO": (Linné, 1758)
cylindricus
sp 990911 ?
In "CGBRO": (Quoy & Gaimard, 1824)
20+, 991105
(116?)
cf extraordinaria. In "CGBRO": (Allan, 1932)
Last OWN Remarks
***
99
990903
991103 991105
990717
990524 990309
990616
990730
990816
990621
990612
990828
990730
990905
990610
First
99
Diaphanidae
Colpodaspis
Siphopteron
Chelidonurid
Facelinidae
Aeolidiidae
Glaucidae
Suborder Aeolidina
Arminidae
Facelinidae
Facelinidae
Facelinidae
Facelinidae
Flabellinidae
Flabellinidae
Flabellinidae
Facelinidae
Facelinidae
Flabellinidae
Glaucidae
Glaucidae
Flabellinidae
Flabellinidae
Flabellinidae
Flabellinidae
Aeolidiidae
Glaucidae
Glaucidae
Glaucidae
Glaucidae
Facelinidae
Glaucidae
Aeolidiidae
Aeolidiidae
Order = Notaspidea
Dermatobranchus
Dermatobranchus
Dermatobranchus
Dermatobranchus
Dermatobranchus
Dermatobranchus
Dermatobranchus
Dermatobranchus
Dermatobranchus
Dermatobranchus
Aeolidiella
Aeolidiella
Cerberilla
Cuthona
Cratena
Cratena
Cuthona
Eubranchus ?
Favorinus
Favorinus
Flabellina
Flabellina
Flabellina
Flabellina
Flabellina
Flabellina
Flabellina
Limenandra
Moridilla
Phidiana
Phyllodesmium
Phyllodesmium
Phyllodesmium
Phyllodesmium
Protaeolidiella
Pteraeolidia
Aeolid
Aeolid
Aeolid
Aeolid
Aeolid
Aeolid
Aeolid
Pleurobranchidae Pleurobranchidae Berthella
Pleurobranchidae Pleurobranchidae Pleurobranchus
Pleurobranchidae Pleurobranchus
Pleurobranchidae Pleurobranchidae Pleurobranchus
Pleurobranchus
Order = Cephalaspidea ?
Diaphanidae
Order = Cephalaspidea
cf. gonatophora
sp
sp
sp
sp
sp
sp
sp
sp
sp
alba
sp
annulata
cf annulata
lineata
sp
sp.
sp
japonicus
tsuruganus
bicolor
delicata
exoptata
ornata
riwo
rubrolineata
sp. 1.
nodosa
brocki
indica
briareum
longicirrum
sp
sp
atra
ianthina
martensi
forskalii
grandis
peroni
sp.
thompsoni
bruneomarginatum
sp
x
Baba, 1955
(Angas, 1864)
Gosliner & Willan, 1991
(O´Donoghue, 1929)
x
Haefelfinger & Stamm, 1958
Bergh, 1888
(Bergh, 1896)
(Bergh, 1896)
(Bergh, 1905)
Baba, 1949
Baba & Abe, 1964
(Kelaart, 1858)
(Gosliner & Willan, 1991)
Gosliner & Willan, 1991
(Eliot, 1905)
Risbec, 1928
(Pilsbry, 1896)
(Ruppel & Leuckart, 1828)
Pease, 1868
Cuvier, 1804
Brown, 1979
1
3
8
3
1
1
1
10
4
1
1
1
3
13
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
2
1
1
3
20
2
19
25
16
27
1
3
1
1
3
1
1
1
3
1
17
398
1
3
4
1
2
1
1
1
3
11
5
1
1
2
1
1
4
1
2
1
1
1
42
4
20
16
1
1
2
2
316
Appendix I, 2(6)
981113
970920
980907
970920
981104
970920
981202
981202
981112
981124
980325
981019
980916
970920
981114
980315
981027
981027
980325
980908
981119
970920
970920
980620
981117
990514
990621
990805
991022
990624
990515 990917
990603 990615
991111
990617
990630
990630
990617
990816
990505 991111
990515 991116
990517 991121
990721
990623 990816
990617 991115
990609
990605
990906
990605 991117
990818
990811
990811 990926
990816
990629 990826
990803
990708 990725
990806
990511
990925
990709 991023
990613 991117
990527 991120
990824
990525
990603 990622
990621
990511
51
101
206
68
90
119
165
198
82
83
117
126
127
129
174
124
85
65
118
148
67
77
6
61
160
53
165
87
192
7
173
96
107
88
cf OWN101
D. gonatophora in "D" 1996 p 295.
similar to OWN 85
(Caloria)
(briareus)
& OWN 148
In Gosliner, 1987 (former Coryphella sp)
(japonicus?)
annulata ?
C. annulata in "G" species 232. (Baba, 1949)
In "GBRO": (Eliot, 1904)
(forskali) In "GBRO": Rüppell & Leuckart, 1828
Buoy 3 -15 specimens
Cephalaspidea in SSF but "unresolved" in CL
In "WB" species 18. Carlsson & Hoff, 1974
Philinopsis/Chelidonura?
Tritoniidae
Tethydidae
Scyllaeidae
Chromodorididae
Chromodorididae
Chromodorididae
Chromodorididae
Chromodorididae
Chromodorididae
Chromodorididae
Chromodorididae
Chromodorididae
Dorididae
Dendrodorididae Dendodorididae
Chromodorididae Chromodorididae
Chromodorididae
Chromodorididae
Dendrodorididae
Dendrodorididae Dendodorididae
Chromodorididae
Chromodorididae
Chromodorididae
Chromodorididae
Chromodorididae
Chromodorididae
Chromodorididae
Chromodorididae
Chromodorididae
Chromodorididae
Chromodorididae
Chromodorididae
Chromodorididae
Suborder Dendronotina
Dorididae
Chromodorididae
Chromodorididae
Chromodorididae
Scyllaeidae
Tethydidae
Arminidae
Suborder Arminina
Bornellidae
Bornellidae
Bornellidae
Bornellidae
Dendronotidae
Tritonidae
Ardeadoris
Asteronotus
Ceratosoma
Ceratosoma
Ceratosoma
Ceratosoma
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Chromodoris
Dendrodoris
Dendrodoris
Dendrodoris
Dendrodoris
Dendrodoris ?
Dendrodoris ?
Diaphorodoris
Discodoris
Bornella
Bornella
Dendronotus
Marionia
Marionia
Melibe
Melibe
Scyllaea
Tritonia
Tritonia
Tritoniopsis
(Tritonopsilla
(Baba, 1938)
Bergh, 1877
mitsui
boholiensis
Rudman, 1982
(Crosse, 1875)
Rudman, 1982
(Stimpson, 1855)
(Kelaart, 1858)
Rudman, 1982
Pruvot-Fol, 1930
Rudman, 1987
Rudman, 1982
Bergh, 1877
Rudman, 1982
(Risbec, 1956)
Gosliner & Behrens, 1998
Bergh, 1877
(Kelaart, 1858)
Risbec, 1928
Rudman, 1984
(Hasselt, 1824)
(Bergh, 1875)
(van Hasselt, 1824)
Abraham, 1876
(J. E. Gray, 1827)
(Garret, 1879)
2
2
1
1
1
4
2
63
1
1
2
3
1
1
1
5
2
106
1
1
1
1
1
1
17
11
81
1
71
1
1
22
13
132
1
119
1
1
1
1
109
6
30
60
8
3
40
(Baba, 1949)
1
1
215
8
35
106
9
3
49
1
1
1
1
7
1
2
1
1
2
1
2
1
2
1
3
1
Risbec, 1937
Alder & Hancock, 1864
(Linneaeus, 1758)
8
1
2
1
1
2
5
3
Johnson, 1983
(Adams & Reeve, 1848)
x
x
2
33
3
x
egretta
cespitosus
miamirana
sinuata
tenue
trilobatum
albopunctata
alius
annae
boucheti
coi
dianae
elisabethina
fidelis
geometrica
hintuanensis
kuiteri
kuniei
leopardus
lochi
cf. lochi
magnifica
preciosa
quadricolour
reticulata
sp
sp
sp
sp
sp
strigata
verrieri
willani
carbuncolosa
nigra
cf. nigra
tuberculosa
anguilla
stellifer
sp
sp.
viridescens
engeli
fimbriata
pelagica
sp.
sp.
alba
australis)
Dermatobranchus sp. 2.
Appendix I, 3(6)
981119
980404
981216
980801
980318
980831
980401
981126
980801
980315
970920
970920
970920
970920
970920
970920
970920
980501
970920
970920
970920
980901
980501
970920
981103
981121
980325
970920
980325
980501
970920
981124
991117
991109
991104
991117
991117
991001
991105
991117
990910
990722
991120
990728
991101
991018
990603
990715
990724
990802 991117
990704 990730
990505 991116
990906
991101 991111
990711 991007
990522
990518
990609
990505
990623
990520
990704
990731
990914
990505
990703
990524
990514
990524
990521
990603
990603 991120
990621
990717 ?
991105
990824
990708
990816
990607 990703
991102
990903 990915
990730
990515 990517
991010 991124
990610 990825
86
201
69
196
179
142
81
100
145
149
2
48
123
157
146
12
137
203
168
131
74
In Gosliner,1987 (former D. Sp)
(Dendrois)
verrieri? albonares
In Debelius 1996 p 200
In "GBW" species 577 (Quoy & Gaimard, 1832) (=tinctoria)
kuiteri ? OS 0184
(elizabethina)
(bouchetti)
In "WB" species 147. Rudman 1987
In "GBRO": (Gray, 1827)
Earlier: Miamira sinuata
(Tritonopsilla)
Probably a misstake!
In "GBRO": Linne, 1758
OWN13/1
In "GBRO": (A. Adams & Reeve in A. Adams, 1848)
Aegiridae
Aegiridae
Polyceridae
Polyceridae
Polyceridae
Phyllidiidae
Phyllidiidae
Phyllidiidae
Phyllidiidae
Chromodorididae
Chromodorididae
Goniodorididae
Polyceridae
Aegiritidae
Aegiritidae
Polyceridae
Polyceridae
Polyceridae
Polyceridae
Polyceridae
Chromodorididae
Chromodorididae
Dorididae
Dorididae
Chromodorididae
Chromodorididae
Chromodorididae
Hexabranchidae Hexabranchidae
Chromodorididae
Gymnodorididae
Dorididae
Dorididae
Dorididae
Gymnodorididae
Gymnodorididae Gymnodorididae
Dorididae
Dorididae
Chromodorididae
Chromodorididae
Discodoris?
Doris?
Durvilledoris
Durvilledoris
Fryeria
Fryeria
Glossodoris
Glossodoris
Glossodoris
Glossodoris
Gymnodoris
Gymnodoris
Gymnodoris
Gymnodoris
Gymnodoris
Gymnodoris
Gymnodoris
Gymnodoris
Gymnodoris
Gymnodoris
Halgerda
Halgerda
Halgerda
Halgerda
Hexabranchus
Hypselodoris
Hypselodoris
Hypselodoris
Hypselodoris
Hypselodoris
Hypselodoris
Hypselodoris
Hypselodoris
Hypselodoris
Hypselodoris
Hypselodoris
Jorunna
Jorunna
Nembrotha
Nembrotha
Nembrotha
Nembrotha
Nembrotha
Nembrotha
Nembrotha
Notodoris
Notodoris
Notodoris
Noumea
Noumea
Noumea
Okenia
Pectenodoris
Pectenodoris
Phyllidia
Phyllidia
liturata
sp. 10.
pusilla
similaris
menindie
rueppelii
albocincta
cincta
hikuerensis
rufomarginata
alba
ceylonica
sp.
sp.
sp.
sp.
sp.
sp.
sp.
sp. 1.
carlsoni / aff. carlsoni
malesso
cf. malesso
sp
sanguineus
bullocki
emma / emmae
infucata
krakatoa
maculosa
cf. maculosa
purpureomaculosa
rudmani
sp.
sp. 1.
whitei
funebris
sp.
chamberlaini
cristata
kubaryana
lineolata
rutilans
sp.
sp. 4.
gardineri
minor
serenae
crocea
varians
sp. 2.
sp.
aurora
trilineata
babai
coelestis
(A. Adams & Reeve 1850)
Brunckhorst, 1993
Bergh, 1905
Rudman, 1986
(Pease, 1871)
x
x
x
Eliot, 1903
Eliot, 1904
Gosliner & Behrens, 1997
Bergh, 1877
Bergh, 1877
Bergh, 1905
(Pruvot-Fol, 1931)
x
(Adams & Reeve, 1850)
(Kelaart, 1858)
(Ruppell & Leuckart, 1830)
(Collingwood, 1881)
Rudman, 1977
(Ruppel & Leuckart, 1828)
Gosliner & Johnsson, 1999
(Pease, 1871)
x
Hamatani, 1995
x
x
Rudman, 1978
Carlson & Hoff, 1993
(Bergh, 1888)
(Pruvot-Fol, 1954)
(Bergh, 1890)
(Bergh, 1877)
(Kelaart, 1858)
Bergh, 1905
x
x
x
Brunckhorst, 1993
Bergh, 1869
1
1
5
1
1
1
1
1
1
5
1
1
1
1
2
2
82
2
3
113
1
1
2
1
10
36
4
2
1
26
1
5
13
5
18
1
29
1
1
2
1
1
3
1
1
1
1
1
1
1
1
5
1
1
1
1
1
1
1
1
2
1
2
1
2
1
1
7
1
2
2
2
1
1
8
1
1
2
Appendix I, 4(6)
970920
981029
980706
970920
970920
970920
970920
970920
970920
970920
970920
981124
980915
980324
970920
970920
970920
981103
980421
970920
981001
970920
970920
981016
981016
970920
980620
980620
980620
981024
981216
970920
980815
991026
991111
991117
991116
990606 990806
990727 990802
990514 991117
990816
990630 990730
990606
990525
990524
991015
991109
990810
990513 991118
991026
57
154
990825
990728
990520 990915
990707
990809
990910
990811
64
103
171
128
76
199
134
20
183
170
66
197
135
104
125
133
158
176
186
205
208
58
45
131
147
169
990527 991116
990524 990906
991111
991001 991014
990605
990911
991102 991105
990628
990704
990803
990825
990914
991103
991120
990605 990806
990721
990702 990717
990718
990811
990603 990901
990621
(=Hopkinsia)
(Mike Millers Slug Site)
& OWN103, OW
varians / norba
In "GBW" species 610. (Gosliner & Behrens 1998)
whitei ex mouaci, OS 0623
In ZJLS 125:1-114
In "GBRO": (Rüppell & Leuckart, 1830)
small pink variety + purple and white variety
emma
carlsoni
as in Debelius 1996 =Halgerda sp. 1.
In "SSF" author not given.
(meinindae)
(cf Fryeria ruppeli)
(Discodoris ? liturata)
Doris sp. 10
Phyllidiidae
Phyllidiidae
Phyllidiidae
Phyllidiidae
Phyllidiidae
Phyllidiidae
Phyllidia
Phyllidia
Phyllidia
Phyllidia
Phyllidia
Phyllidia
Phyllidia
Phyllidia
Phyllidia
Phyllidia
Phyllidia
Phyllidia
Phyllidia
Phyllidiidae
Phyllidia
Phyllidiidae
Phyllidiidae
Phyllidia
Phyllidiidae
Phyllidiella
Phyllidiidae
Phyllidiidae
Phyllidiella
Phyllidiidae
Phyllidiidae
Phyllidiella
Phyllidiidae
Phyllidiidae
Phyllidiella
Phyllidiidae
Phyllidiopsis
Phyllidiidae
Phyllidiopsis
Phyllidiidae
Phyllidiopsis
Phyllidiidae
Phyllidiopsis
Phyllidiidae
Phyllidiidae
Phyllidiopsis
Phyllidiidae
Phyllidiidae
Phyllidiopsis
Phyllidiidae
Phyllidiidae
Phyllidiopsis
Phyllidiidae
Phyllidiopsis
Phyllidiidae
Phyllidiidae
Platydoris
Dorididae
Platydoris
Platydoris
Plocamopherus
Triophidae
Polyceridae
Polycera
Polyceridae
Polycera ?
Reticulidia
Phyllidiidae
Phyllidiidae
Reticulidia
Phyllidiidae
Phyllidiidae
Risbecia
Chromodorididae
Chromodorididae Chromodorididae Risbecia
Roboastra
Polyceridae
Sebadoris
Dorididae
Tambja
Polyceridae
Polyceridae
Tambja
Polyceridae
Thecacera
Polyceridae
Thecacera
Polyceridae
Polyceridae
Thecacera
Polyceridae
Thorunna
Chromodorididae
Chromodorididae Chromodorididae Thorunna
Trapania
Trapania
Goniodorididae
Trapania
Goniodorididae Gonodorididae
Trapania ?
Trapania ?
Trippa
Dorididae
Dorididae
Trippa
Trippa
Chromodorid
Chromodorid
intecta
sp
sp
fungia
halgerda
imperialis
tryoni
gracilis
nubilosa
morosa
sp.
pacifica
picta
sp
australis
furtiva
sp.
sp. 2.
toddi
sp. cf. coelestis
elegans
exquisita
ocellata
sp
sp
sp
sp
sp
sp
sp
sp
tula
varicosa
willani
nigra
pustulosa
rudmani
zeylanica
annae
dautzenbergi
fissuratus
krempfi
pipeki
shireenae
sphingis
striata
formosa
scabra
sp
ceylonica / ceylonicus
sp. 3.
(Kelaart, 1858b)
x
Rudman, 1987
Brunckhorst & Gosliner, 1993
Brunckhorst & Burn, 1990
(Pease, 1860)
(Garrett, 1873)
(Bergh, 1877)
Pease, 1891
(Bergh, 1877)
x
(Bergh, 1883)
Baba, 1972
x
(Risbec, 1928)
Bergh, 1878
(Kelaart, 1858)
x
Marcus & Marcus, 1970
Lamarck, 1801
Brunckhorst, 1993
(van Hasselt, 1824)
(Cuvier, 1804)
Brunckhorst, 1993
(Kelaart, 1859)
Brunckhorst, 1993
(Vayssiere, 1911)
Brunckhorst, 1993
Pruvot-Fol, 1957
Brunckhorst, 1993
Brunckhorst, 1993
Brunckhorst, 1993
Bergh, 1888
(Alder & Hancock, 1864)
(Cuvier, 1804)
Bergh, 1869
Brunckhorst, 1993
Cuvier, 1804
8
13
1
2
2
1
7
16
1
9
16
1
1
1
1
2
2
14
1
15
1
1
1
1
2
3
1
115
8
19
3
1
1
6
1
1
1
1
1
96
6
4
289
24
8
2
1
1
2
52
11
1
28
2
4
1
2
1
1
164
9
19
3
1
1
6
1
1
1
1
1
131
7
15
604
30
10
2
1
1
2
58
12
1
34
5
7
2
3
1
Appendix I, 5(6)
980530
980624
980909
970920
970920
970920
970920
970920
981121
981119
980526
981124
981202
970920
970920
980919
980319
970920
970920
970920
970920
970920
981017
980901
970920
981002
970920
970920
970920
990925
991118
991103
991027
991101
991124
991113
991117
991109
990917
991124
991028
991112
990805
991115
991117
991105
991105
990812
990515
990617
990704
990728
990613 991116
990614 990903
990603 991116
991022 991111
990609 991113
990515 991112
990521
990619 991118
990609
990807
990519
990721
990624
990709
990623
990708
990712
990709
990713
990714
990729
990707
990515
990628
990515
990505
990619
990622
990802
991018
990930
990917
990519
990626
990828
990604
990520
990716
990820
990906
990920
112
34
39
91
132
151
71
30
75
94
190
181
188
84
195
193
185
59
140
130
136
138
139
143
144
153
167
In "SSF" author not given.
In "GBRO": Brunckhorst & Gosliner in Brunckhorst, 1993
In "GBRO": Brunckhorst & Gosliner in Brunckhorst, 1993
ceylonicus
(shirinae)
In "GBRO": (Hasselt, 1824). Lahoa Channel
990708 + 1 ?
3523
2
4
2
25
1
11
3
3
4
2
1
4
49
5
36
1
13
5
4
4
2
1
1
1
1
1
12
2
2
1
1
15
3
2
1
1
1
1
2
4
1
1
1
3
5
1
* = Family designation according to "Sea Slug Forum"
**= Family designation according to "Checklist of the Great Barrier Reef Ophistobranchia" (in M & W, 1999)
***= Operation Wallacea Nudibranchia (Internal number)
(Trinchese, 1893)
(Marcus, 1965)
Gosliner, 1995
Gosliner, 1995
x
x
(Montagu, 1804)
Bergh, 1872
(Swainson, 1840)
x
x
x
Bergh, 1870
2888
159
Plakobranchiidae
Plakobranchiidae
Plakobranchiidae
Plakobranchiidae
Plakobranchiidae
Polybranchiidae
sp
sp. 1.
sp. 2
elegans
brycei
ornata
pilosa ?
sp
sp
sp
sp
sp
sp.
sp.
sp. 1.
sp. 8.
viridis
pusilla
sp
cremoniana
bayeri
carlsoni
hoffae
kathae
lineolata
sp
sp
undula
Order = Sacoglossa
Elysiidae
Limapontiidae
Elysiidae
Elysiidae
Elysiidae
Elysiidae
Elysiidae
Elysiidae
Elysiidae
Costasiellidae
Costasiellidae
Caliphyllidae
Costasiella
Costasiella
Costasiella ?
Cyerce
Elysia
Elysia
Elysia
Elysia
Elysia
Elysia
Elysia
Elysia
Elysia
Elysia
Elysia
Elysia
Elysia
Elysiella
Plakobranchus
Stiliger
Thuridilla
Thuridilla
Thuridilla
Thuridilla
Thuridilla
Thuridilla
Thuridilla
Thuridilla
Order = Nudibranchia
(Aeolidina = 483, Arminina = 30, Dendronotina = 47, Doridina = 2328)
Suborder Doridina
Appendix I, 6(6)
981116
980315
980907
981024
980507
980620
980825
980620
981008
981005
980801
981019
981029
980621
981124
990609
990524
990925
990515
990910
990603
990721
990512
990517
990609
990805
990906
990807
990917
990824
990914
990621
990610
990701
991110
990925
991115
990804
990515 991117
99064 990615
990616 990617
990728
990518
991117
990716 991101
990617 990910
990621
92
109
25
47
191
62
63
99
105
111
150
54
159
55
180
120
80
93
207
In "GBW" species 550. Gosliner 1995
In "D" 1996 p 169.
hoffae (livida ?)
In "GBRO": (Er. Marcus, 1965)
Nothern species!?
cf filicauda
In "GBRO": Bergh, 1871
In "WB" species 70. (Jensen & Wells, 1990) + "D" 1996
Date
991110
Species:
Chromodoris annae
Jorunna funebris
Jorunna funebris
Phyllidia varicosa
Jorunna funebris
Date
991111
Species:
Chromodoris annae
Chomodoris elisabethina
Phyllidia varicosa
Phyllidia coelestis
Date
991112
Species:
Chromodoris annae
Chromodoris annae
Chromodoris dianae
Date
991108
Species:
Jorunna funebris
Jorunna funebris
Date
991109
Species:
Nembrotha kubaryana
Jorunna funebris
Jorunna funebris
Phyllidia varicosa
Divers
LK/PB/JH
Size (mm)
25
50
20
30
Divers
LK/PB
Size (mm)
56
35
20
28
35
Buoy 3 "+3m" (Latitude: S 05 28 386, Longitude: E 123 45 439) Searcharea 150 sqm
Afternoon dives ("PM")
Morning dives ("AM")
Searchtime (min) Line depth (m)
Divers
Date
4,6
24
RS/NR/SY
991108
4,5
20
V-C (cm)
H-C (cm)
Depth (m)
Size (mm)
Species:
Colour
Surface
V-C (cm)
H-C (cm)
Depth (m)
64
190
7
40
Chromodoris annae
CA
290
300
6,6
64
190
7
36
Chromodoris annae
DCA
230
430
6,1
95
680
5
20
SPL
-190
1480
4,6
95
680
5
20
SPL
-190
1480
4,6
Divers
Date
4,5
24
LK/PB
991109
4,7
27
V-C (cm)
H-C (cm)
Depth (m)
Size (mm)
Species:
Colour
Surface
V-C (cm)
H-C (cm)
Depth (m)
5
190
6,4
25
Chromodoris annae
Sea cucumber!
100
1080
5,8
5
190
6,4
25
Chromodoris annae
SP
-50
1490
5,2
1
200
7,2
60
SP
-50
1490
5,2
10
420
4,2
40
Brown
CE
56
1640
6,1
11
1485
5,1
30
Jorunna funebris
Pink
CA
-56
2460
4,8
11
1485
5,1
20
Jorunna funebris
Divers
Date
Divers
4,9
27
RS/JH/SY/EW
991110
40
RS/JH/SY/EW
Marker
H-C (cm)
Depth (m)
Size (mm)
Distance (cm) Species:
Colour
Surface
Marker
H-C (cm)
Depth (m)
Size (mm)
10
420
4,2
35
18 NE
Pink
CA
18
130
5,9
30
20
750
5,4
25
22 N
DC
17
130
5,8
40
11
1485
5,4
30
Jorunna funebris
35 SW
10
420
4,2
35
11
1485
5,4
20
Jorunna funebris
7 SW
Orange
SPS
20
750
5,1
25
23
2250
6,4
30
varicosa
Phyllidia
NW
26
Brown
CE
11
1485
5,1
30
26 NW
Brown
CE
11
1485
5,1
20
8N
S
23
2250
5,5
30
9N
RCK
22
2950
4,5
60
Divers
Date
Divers
4,8
28
LK/PB/GY/TG
991111
3,8
20
LK/PB/GY/TG
Marker
H-C (cm)
Depth (m)
Size (mm)
Colour
Surface
Marker
H-C (cm)
Depth (m)
Size (mm)
32
200
5
35
10 NE
DC
73
6
7,1
40
12
950
5
70 NE
MA
17
110
4,9
30
2
2060
45
Jorunna funebris
37 NW
80
380
3,1
30
60 NW
69
670
3,8
45
9 NW
23
2250
8E
DC
23
2300
5
30
10 SW
36
2980
6,4
60
Divers
Date
Divers
4,9
32
RS/JH/SY/EW
991112
5,7
28
RS/JH/SY/EW
Marker
H-C (cm)
Depth (m)
Size (mm)
Colour
Surface
Marker
H-C (cm)
Depth (m)
Size (mm)
5
7,6
40
Chromodoris annae
42 E
Brown
MA
73
0
7,2
40
5
8,3
43
Chromodoris annae
230 SW
Brown
CS
5
240
6,7
40
80
4,4
15
67 S
Grey
SPG
10
440
3,8
10
4,4
44
66 N
SP
72
670
4,6
28
72/69
5,2
28
20 W
DC
20
690
4,7
30
17
6,7
40
Chromodoris annae
15 NE
MA
36
5
35
18
6,7
40
Chromodoris annae
8N
23
38
73
8
25
Chromodoris annae
36
Chromodoris dianae
Appendix II, 1(4)
Search records from "Spatial distribution of nudibranchs according to benthic structure"
Surface Colour Distance (cm)
220 W
CS
214 SW
DC
90 W
CT
62 E
CT
94 NE
CA
130 E
TA
130 E
TA
80 E
Brown
CS
70 NE
10 N
DC
10 NE
R
0
CE
40 W
SPT
29 SE
OT
26 NW
Brown
CE
26 NW
Brown
CE
7 NE
0
AA
0
AA
0
SP
5N
Shell
9N
SP
9N
SP
Surface Colour
SPS Orange
SPS Orange
Grey
SPE
Grey
SPE
Divers
LK/PB
Size (mm)
60
25
50
24
15
Divers
RS/JH/SY/EW
Size (mm)
35
20
17
30
45
40
Divers
LK/PB/GY/TG
Size (mm)
20
25
20
Divers
RS/JH/SY/EW
Size (mm)
Date
991109
Species:
Phyllidia varicosa
Phyllidia coelestis
Date
991110
Species:
Nembrotha kubaryana
Date
991111
Species:
Date
991112
Species:
Chromodoris elisbethina
Jorunna funebris
13
26
45
30
Divers
LK/PB/JH
Size (mm)
20
40
25
25
30
40
70
45
Date
991108
Species:
Reticulidia fungia
Jorunna funebris
Nembrotha kubaryana
Chromodoris dianae
Jorunna funebris
Buoy 3 "+12m" (Latitude: S 05 28 386, Longitude: E 123 45 439) Searcharea 150 sqm
Divers
Date
RS/NR/SY
991108
11,8
37
Size (mm)
Species:
Colour
V-C (cm) Surface
H-C (cm)
Depth (m)
30
110
130
13
55
Phyllidia coelestis
Brown
MA
-315
532
9,3
13
MA
-33
765
11,9
20
Reticulidia fungia
Purple
SPE
-190
1000
10,3
5
Reticulidia fungia
CA
160
1050
13,5
20
Nembrotha kubaryana
Line
0
1105
11,9
25
CA
40
1860
11,7
Nembrotha kubaryana
Pink
CA
-325
2060
9
20
Divers
Date
LK/PB
991109
12,2
35
Size (mm)
Species:
Colour
V-C (cm) Surface
H-C (cm)
Depth (m)
25
Grey/Green
AE
250
450
15,8
35
Purple
TA
-190
880
10,9
50
CA
-120
1800
10,9
15
SPE
65
2300
12,7
RCK
-50
3000
10,4
Divers
Date
RS/JH/SY/EW
991110
40
Size (mm)
Colour
Surface
Marker
H-C (cm)
Depth (m)
25
kubaryana
Nembrotha
SE
11
AA
27
135
13,5
30
6 SW
CA
19
170
14
40
7W
RCK
9
300
11
13 E
Purple
SPE
7
900
11
4W
Grey
SPE
16
1345
12,6
90 E
RCK
6
1795
13
Divers
Date
LK/PB/GY/TG
991111
11,7
15
Size (mm)
Colour
Surface
Marker
H-C (cm)
Depth (m)
20
14 NE
AA
33
0
10.0
15
5 SW
CA
19
170
12,7
20
Chromodoris annae
30 N
SPE
16
1345
12,1
40
70
Jorunna funebris
15
Divers
Date
RS/JH/SY/EW
991112
12
21
Size (mm)
Colour
Surface
Marker
H-C (cm)
Depth (m)
28
6 SE
CA
19
170
12,8
56 SE
DC
16
1346
12,2
Chromodoris annae
42 SE
CA
34
1580
13
85 NE
AA
37
2400
12
Jorunna funebris
217 E
MA
6
1800
11
32
Chromodoris dianae
30
Appendix II, 2(4)
14
12,7
22
H-C (cm)
Depth (m)
170
10
900
11
1340
11
1640
12
2380
12,5
2880
12
13
26
H-C (cm)
Depth (m)
170
13,8
900
11,4
13,2
1615
13,4
Marker
19
7
16
34
37
3
6
Marker
19
7
16
17
27
35
Dark
Surface Colour
DC
SPE
Yellow
HY
SPG
SPM
SP
CA
SPE
Surface Colour
Distance (cm)
50 SW
35 W
69 SE
59 SE
189 NE
268 NE
230 E
13 N
Distance (cm)
1 SE
1E
18 N
1S
Distance (cm)
11 SE
10 NE
95 E
Surface Colour
AA
SPE
RCK
13
28
H-C (cm)
Depth (m)
135
14,2
900
11,5
1795
13
Marker
27
7
6
Distance (cm)
11 NE
0
7 NE
7W
12,1
56
V-C (cm) Surface Colour
H-C (cm)
Depth (m)
RCK
80
175
13,5
Purple
CA
137
340
15,6
RCK
74
460
13,8
120
560
12,5
120
560
12,5
SC
-40
885
12,8
Purple
SPE
-150
1000
11,3
Red
TA
10
2980
11,3
Pink
CA
-80
3000
10,4
12
25
H-C (cm)
Depth (m)
CA
7
900
11
Red
MA
6
1800
13,5
CA
8
1920
11
CA
9
3000
10,4
Date
991109
Species:
Chromodoris annea
Jorunna funebris
Jorunna funebris
Date
991110
Species:
Phyllidia varicosa
Jorunna funebris
Jorunna funebris
?
Date
991111
Species:
Phyllidia coelestis
Phyllidia coelestis
Phyllidia varicosa
Chromodoris annea
Jorunna funebris
Jorunna funebris
Date
991112
Species:
Chromodoris annea
Jorunna funebris
Jorunna funebris
Phyllidia coelestis
Date
991108
Species:
Chromodoris annea
Jorunna funebris
Jorunna funebris
5,3
5,6
5,6
6,5
20
25
25
15
5,7
Searchtime (min)
25
Depth (m)
6
Searchtime (min)
28
Depth (m)
6,8
7,2
7,7
7,7
7,5
7,8
7,5
Searchtime (min)
28
Depth (m)
7,7
8,1
7,3
7,3
7
6,7
Searchtime (min)
40
Depth (m)
4,4
5,7
7,9
6,5
8
6,3
7,9
620
1000
39
46
V-C (cm)
H-C (cm)
-75
310
-32
1220
100
2450
100
2450
75
2880
-40
2920
50
2965
Line depth (m)
5,3
Marker
H-C (cm)
46
1000
47
2180
48
2470
48
2470
38
2680
49
2940
Line depth (m)
5,6
Marker
H-C (cm)
60
110
39
620
46
1000
43
1177
47
2180
41
2360
24
2460
48
2470
48
2470
74
2720
49
2940
49?
3000
49?
3000
Line depth (m)
5,2
Marker
H-C (cm)
43
1130
24
2530
74
2680
42
3000
42
3000
48
2470
Line depth (m)
5,7
39
V-C (cm)
H-C (cm)
Depth (m)
-180
150
3,9
-120
330
4,8
-240
900
4,3
-40
1220
5,6
0
1228
6
45
2415
8
70
2415
8,3
70
2415
8,3
Divers
RS/JH/SY
Size (mm)
20
15
50
40
26
25
40
Divers
LK/PB/GY
Size (mm)
35
25
70
40
25
25
Divers
RS/JH/SY
Size (mm)
20
11
40
11
25
30
40
70
40
30
25
60
22
Divers
LK/PB/GY/TG
Size (mm)
25
Divers
RS/NR/SY
Size (mm)
10
25
25
20
20
34
70
50
CA
CA
Surface
SPE
SPE
SPE
RCK
RCK
Colour Distance (cm)
60 SW
10 E
7 NW
8 SE
8 SE
105 NW
40 W
40 SE
95 W
40 NE
24 S
38 W
11 SW
10 SE
4W
Brown
18 E
53 SW
42 NW
12 SW
27 S
8 SE
8 SE
Surface
OT
SP
AA
MA
SPE
CM
5S
Grey
7 SE
Yellow
41 NW
28 NE
?
12 S
Pink
Orange
SPS
SP
Surface
SPE
SPM
CM
CA
DC
CA
Colour
Yellow
Red
Red
Pink
Pink
Grey
Colour
Surface
SPE
AE
CM
CM
Surface
R
R
CA
SPB
SPE
RCK
SPE
SPE
Date
991112
Species:
Chromodoris annea
Jorunna funebris
Jorunna funebris
Date
991111
Species:
Phyllidia coelestis
Phyllidia varicosa
Chromodoris annea
Jorunna funebris
Jorunna funebris
Thecacera picta
Date
991110
Species:
Phyllidia coelestis
Phyllidia varicosa
Ptereoalidia ianthina
Date
991108
Species:
Phyllidia varicosa
Chromodoris annea
Jorunna funebris
Jorunna funebris
Date
991109
Species:
Phyllidia varicosa
Jorunna funebris
Jorunna funebris
7,3
30
45
8
8
Searchtime (min)
Divers
23
LK/PB/GY/TG
Depth (m)
Size (mm)
7
20
6,9
25
6,9
20
8
35
Marker
43
42
42
24
47
48
48
74
46
6,5
6,9
11
30
Line depth (m)
6,8
H-C (cm)
1130
3000
3000
2530
2180
2530
2530
2720
Marker
39
46
43
74
47
41
24
48
48
42
42
Line depth (m)
6,8
H-C (cm)
610
1000
1177
2120
2180
2360
2460
2470
2470
3000
3000
Searchtime (min)
31
Depth (m)
6,5
Divers
RS/JH/SY
Size (mm)
22
16
To "laboratory"
Marker
39
46
51
26
Line depth (m)
6,8
H-C (cm)
630
1000
1000
1200
60 SE
Grey
SPE
6E
CA
6E
CA
31 SE
DC
16 E
CA
40 W
40 W
8 NW
CA
97 W
CE
24 S
SP
50 W
11 SW
White
SPE
36 NE
SPE Orange
15 W
16 E
CE
18 E
18 N
36 NW
10 E
10 E
SP
27 S
26 NW
CA
14 SW
Black
SPD
14 SW
Grey
SPE
6,9
37
H-C (cm)
Depth (m)
DC
-186
80
6,3
Pink
CA
-100
345
6,6
Grey
SPE
-31
1245
7,1
Brown
CA
140
2390
9
Green
CE
100
2480
8,7
Grey
SPE
100
2480
8,7
CA
-50
2910
7,1
Pink
CA
80
2950
8,6
Pink
CA
80
2950
8,6
25
H-C (cm)
Depth (m)
RCK
170
950
9,1
RCK
140
1970
8,6
CM
100
2450
7,7
CM
100
2450
7,7
Searchtime (min)
Divers
23
LK/PB/GY/TG
Depth (m)
Size (mm)
6,3
18
9
35
9,6
30
6,1
35
Divers
LK/PB/JH
Size (mm)
20
20
15
35
55
40
25
35
30
Divers
RS/JH/SY
Size (mm)
40
15
50
40
Sampela "+3m" (Latitude S 05 29 041, Longitude E 123 45 271) Searcharea 150 sqm
Appendix II, 3(4)
Searchtime (min)
49
Depth (m)
12,8
13,1
13,3
11,8
13,1
14,4
14
12,6
12,6
Searchtime (min)
16
Depth (m)
11,9
Divers
RS/JH/SY
Size (mm)
33
25
26
30
11
30
26
30
18
Divers
LK/PB/GY
Size (mm)
35
Date
991109
Species:
Phyllidia sp
Phyllidia coelestis
Phyllidia coelestis
Phyllidia coelestis
Chromodoris coi
Date
991110
Species:
Date
991111
Species:
Phyllidia coelestis
P.s ?
Chromodoris coi
Date
991112
Species:
Chromodoris coi
Jorunna funebris
Jorunna funebris
10
20
11
12,3
Searchtime (min)
Divers
30
RS/JH/SY
Depth (m)
Size (mm)
10,6
12
14
12
11,9
30
11,4
15
13,3
50
12,4
30
Searchtime (min)
Divers
28
LK/PB/GY/TG
Depth (m)
Size (mm)
10
35
Searchtime (min)
44
Depth (m)
11,9
13,3
11,6
14,2
10
14
12,8
Divers
RS/NR/SY
Size (mm)
14
40
30
17
40
12
28
Date
991108
Species:
Phyllidia coelestis
Phyllidia varicosa
Sampela "+12m" (Latitude S 05 29 041, Longitude E 123 45 271) Searcharea 150 sqm
Divers
Date
Line depth (m)
LK/PB/JH
991108
12,7
Size (mm)
Species:
Colour
V-C (cm) Surface
H-C (cm)
30
S
10
250
30
CA
167
325
30
Yellow
SPD
-85
642
35
Phyllidia coelestis
SPM Brown/yellow
110
670
10
Pink
CA
-270
1270
25
Phyllidia varicosa
Purple
CA
170
1470
20
Pink
CA
0
2450
20
Divers
Date
Line depth (m)
RS/JH/SY
991109
Size (mm)
Species:
Colour
V-C (cm) Surface
H-C (cm)
26
Phyllidia sp
S
42
0
40
Phyllidia varicosa
RCK
80
280
25
AE
90
410
30
Dark red
SPE
-160
620
30
Phyllidia coelestis
Brown
SP
93
710
11
Phyllidia coelestis
S
-130
1695
12
S
133
2255
Orange
SPS
-130
2422
Pink
CA
-290
2820
Divers
Date
Line depth (m)
LK/PB/GY/TG
991110
12,2
Size (mm)
Colour
Marker Surface
H-C (cm)
No Opisthobranchs found
3N
CA
40
2450
26 S
31
2470
Divers
Date
Line depth (m)
RS/JH/SY
991111
12,1
Size (mm)
Colour
Marker Surface
H-C (cm)
12
3E
Orange
SPS
52
240
25
13
Yellow
SP
44
690
12
Phyllidia coelestis
60 S
DC
67
1190
8
Jorunna funebris
12 N
Orange
SPE
53
1325
18
Phyllidia varicosa
10 NE
AA
25
1620
22
60 E
OT
31
2430
Divers
Date
Line depth (m)
LK/PB/GY/TG
991112
11,7
Size (mm)
Colour
Marker Surface
H-C (cm)
18
Phyllidia coelestis
100 N
Orange
CA
25
1530
35
Chromodoris coi
12
35
Chromodoris coi
12
60
Chromodoris kunei
44 NW
AE
53
1200
50
Chromodoris kunei
6N
Red
SPE
31/40
2420
10
25
Appendix II, 4(4)
13,4
32
H-C (cm)
Depth (m)
240
12,1
765
13
689
14
1290
14,6
1890
15
2420
15
13
23
H-C (cm)
Depth (m)
690
14,5
1395
12
1620
12
1890
13,5
1890
13,5
1325
12,8
2420
13,5
13,3
16
H-C (cm)
Depth (m)
Marker
44
12?
25
68
68
53
31/40
Marker
52
65
44
12
68
31
Marker
Distance (cm)
12 SW
76 S of tape
200 NE
40 SE
40 SE
39 NW
5N
Yellow
Colour
Grey
SPB
SPB
MA
OT
Surface
SPD
SPE
CA
CA
Distance (cm)
2 SE
16 SE
13 NE
5E
16 W
8E
Distance (cm)
Colour
Colour
Surface
Orange
Surface
13,6
32
Colour
V-C (cm) Surface
H-C (cm)
Depth (m)
Grey
SPE
30
100
13,7
Brown/Yellow
SPM
110
230
14,8
S
80
310
14,5
Yellow
SPT
-140
625
13,2
CA
150
1410
15,1
DC
-70
1595
14,6
Orange
SPC
-280
1730
11,5
CA
-70
2250
13,3
52
Colour
V-C (cm) Surface
H-C (cm)
Depth (m)
S
30
0
14
RCK
179
150
9,1
R
120
270
13,7
S
80
565
14,5
"Dark"
SPE
-160
620
13,2
Brown
SP
93
710
13,8
DC
-60
1340
14,5

Opisthobranchia

Transcription

Similar documents

Full PDF - Check List: Journal of Species Lists and Distribution

2013 CSTR Annual Report - California Ships to Reefs

Mollusca - Chitons, Snails, Clams, Octopus, and More! Domain

Friend or Foe

Destin Jetty Site Guide

Building a Legacy with New York City`s Harbor School - Eco

November - Bay Area Divers

13-604-1042 CMC SHARC Flyer Web PRs

to see Scubalab BC Tester Choice

snorkeling-and-diving