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R EGU L A R PA P E R
A comparative morphological investigation of otoliths of sixparrotfish species (Scaridae) from the Solomon Islands
Antirostrum Short, broad Long or short, broad Long or short, broad Long, broad
Collum Bridge-like Bridge-like, poorly developed Bridge-like, poorlydeveloped
Bridge-like
Excisura Absent Absent or narrow, shallownotch, narrow angle
Absent Medium, shallow notch,medium angle
Note. LT: Total length.
1052 JAWADFISH
(1995), which suggests that it is impossible to provide a key for the
genera based on the shape of the otoliths for the Scaridae. The ante-
rior margin has an anterodorsal angle, which is high in the scarids. The
anterior margin cannot be described as one unit compared with the
other margins; its different components, rostrum, excisura and anti-
rostrum, were described separately. The three other margins of the
otolith, dorsal, ventral and posterior, show a low range in variation.
There were grades of lobation and irregularities across the different
length groups. Similar findings were reported for triplefin species by
Jawad et al. (2007) and for the greater lizardfish Saurida tumbil (Bloch
1795) by Jawad (2008).
The ostium and cauda in all six species studied are flared shape,
mostly shallow and their floor is covered with either lumps of differ-
ent sizes or ridges. The shape of the sulcus acusticus, the crista supe-
rior and crista inferior is also very variable. Shape and nucleus location
result from the release of soluble Ca2+ on the proximal side (Ibsch
et al., 2004), which in turn precipitates as CaCO3 crystals due to an
increasing alkaline gradient, from the sulcal area towards the otolith
edge (Gauldie & Nelson, 1990). As a result, the growth of the crista
superior and crista inferior is preferred and there is a more important
development on the mesial side. The macula is elongated and narrow
in teleosts and the crista superior and inferior are proportionally more
important than the colliculum (Ladich & Popper, 2001; Lombarte &
Fortuno, 1992; Popper & Hoxter, 1981). The macula faces the collum
and prevents otolith growth at this level (Lombarte & Fortuno, 1992;
Pannella, 1980; Popper & Hoxter, 1981). This is clear in some species
where the collum is either poorly developed or absent. Lombarte et al.
(2003) showed that the variability in the shape of the sagittal otolith is
related to genetic, ontogenetic and environmental factors in Merluc-
cius Rafinesque 1810. Previous studies on fossil and extant otoliths
Crista superior(a)
(b)
Posterior
An�rostrum
Anterrior
1·0 mm
Excisura
5.0kV 15.7 mm x30 (SE(M) 1.00 mm
Sulcus
Ventral
Os�um
Dorsal
Cauda
Crista superior
FIGURE 2 (a) The mesial surface of the left otolith of Hipposcarus
longiceps, 187 mm total length (LT), illustrating various features foundon the otolith and described in the text. (b) the mesial surface of theleft otolith of Scarus oviceps, 230 mm LT, illustrating excisura
(a) (b) (c)
(d) (e) (f)
(g) (h) (i)
(j) (k) 1 mm
FIGURE 3 Otoliths of Hipposcarus longiceps taken from fish of
different total lengths: (a) 187 mm (right-side otolith), (b) 196 mm(right), (c) 209 mm (left), (d) 220 mm (right), (e) 230 mm (left), (f )245 mm (right), (g) 255 mm (right), (h) 266 mm (left), (i) 274 mm (left),(j) 328 mm (right), (k) 360 mm (right)
(a) (b) (c)
(d) (e) (f)
(g) (h) 1 mm
FIGURE 4 Otoliths of Scarus dimidiatus taken from fish of different
total lengths: (a) 114 mm (left-side otolith), (b) 115 mm (left),(c) 150 mm (left), (d) 160 mm (left), (e) 178 mm (left), (f ) 187 mm(right), (g) 195 mm (left), (h) 211 mm (right)
JAWAD 1053FISH
have demonstrated that the sulcus morphology usually is consistent
among the species of a single genus (Nolf, 1985, 2013), which sug-
gests that this feature is probably genetically controlled
(Gauldie, 1988).
Interspecific variation in sulcus morphology has previously been
recorded for only a few other genera. For example, the Euro-African
(a) (b) (c)
(d) (e) (f)
(g) (h) (i)
(j) (k)
(l)1 mm
FIGURE 5 Otoliths of Scarus ghobban taken from fish of different
total lengths: (a) 163 mm (right-side otolith), (b) 185 mm (left),(c) 194 mm (left), (d) 205 mm (right), (e) 215 mm (right), (f ) 227 mm(right), (g) 231 mm (right), (h) 245 mm (right), (i) 271 mm (right),(j) 283 mm (right left), (k) 283 mm (left), (l) 319 mm (right)
(a) (b) (c)
(d) (e) (f)
(g) (h) 1 mm
FIGURE 7 Otoliths of Scarus oviceps taken from fish of different total
lengths: (a) 115 mm (right-side otolith), (b) 133 mm (left), (c) 149 mm(right), (d) 156 mm (right), (e) 208 mm (right), (f ) 214 mm (right),
(g) 215 mm (left), (h) 230 mm (right)
(a) (b) (c)
(d) (e) (f)
(g) (h) (i)
(j) (k) 1 mm
1 mm
FIGURE 8 Otoliths of Scarus rivulatus taken from fish of different
total lengths: (a) 126 mm (right-side otolith), (b) 138 mm (right),(c) 144 mm (right), (d) 155 mm (right), (e) 168 mm (right), (f ) 175 mm(left), (g) 182 mm (left), (h) 191 mm (left), (i) 204 mm (right),(j) 217 mm (left left), (k) 261 mm (right)
(a) (b) (c)
(d) (e) (f)
(g) (h) (i) 1 mm
FIGURE 6 Otoliths of Scarus niger taken from fish of different total
lengths: (a) 140 mm (right-side otolith), (b) 151 mm (left), (c) 161 mm(left), (d) 172 mm (right), (e) 183 mm (right), (f ) 197 mm (left right)(g) 202 mm (left), (h) 211 mm (right), (i) 215 mm (right)
1054 JAWADFISH
TABLE
7Geo
grap
hicalvariationin
themorpho
logy
oftheotolithofHippo
scarus
longiceps,Scarus
ghob
ban,
Scarus
niger,an
dScarus
rivulatus
SolomonIsland
sArabian
–Persian
gulf
SouthAfrica
New
Caled
onia
Taiwan
Present
stud
ySa
digh
zade
h&
Tuset,2
012
Smaleet
al.,1995
Rivatonan
dBourret,1
999
Linan
dChan
g,2012
Hippo
scarus
longiceps
201mm
L T–
––
–203mm
L T
Oval
––
––
Shap
e:Ovate
Coarsely
lobe
d–
––
–Posteriormargin:S
traigh
t
Short
––
––
Rostrum:L
ong
Smooth,p
resenc
eofinde
ntation
––
––
Ostium:F
loorwithco
arse
lumps,noinden
tation
anteriorly
Scarus
ghob
ban
194mm
L T215mm
L T245mm
L T319mm
L T191mm
L T210mm
L T241mm
L T375mm
L T
Oblong
Oblong
Oval
Pointed
Shap
e;cu
neiform
-elliptic
Shap
e:Circu
lar
Shap
e:Oblong
Rostrum:R
ounded
Long
Lobe
dLo
bed
–Rostrum:S
hort
Posteriormargin:
With
notch,
nearly
smooth
Dorsalmargin:E
marginate
–
Absen
tEmargina
teSlightly
emargina
te–
Excisura:
Nono
tch
Ven
tralmargin:
Crena
teVen
tralmargin:L
obed
–
Welld
evelope
dLo
ng,p
ointed
Lobe
d–
Cristasupe
rior
andinferior:
Poorlyde
velope
d
Rostrum:S
hort,b
road
Rostrum:E
marginate
–
–Bridg
e-like
Lobe
d–
–Collu
m:A
bsen
tPosteriormargin:P
ointed
–
–W
elld
evelope
dOstial
––
Cristasupe
rior:Poorlyde
velope
dSu
lcusacusticus:Ostio-cau
dal
–
–Dee
pDev
elope
d–
–Dorsalde
pression:
Shallow
Collu
m:A
bsent
–
–245mm
L TW
elld
evelope
d–
–245mm
L TCristasuperior:Absent
–
–Slightly
emargina
teW
ithridg
es–
–Ven
tralmargin:
Crena
teOstium:S
mooth
–
–Pointed
––
–Rostrum:B
road
––
–Bridg
e-like
––
–Collu
m:A
bsen
t–
–
–W
elld
evelope
d–
––
Cristasupe
rioran
dinferior:
Poorlyde
velope
d–
–
–Dee
p,floorwithridg
es–
––
Cau
da:S
hallo
w,smooth
floor
––
–Dee
p,floorwithridg
es–
––
Ostium:S
hallo
w,floorwithlumps
––
–319mm
––
–430mm
––
–Oblong
––
–Sh
ape:
Circu
lar
––
–Coarse
lobe
d–
––
Dorsalan
dve
ntralm
argins;
fine
cren
ate
––
–Lo
bed,
long
––
–Rostrum:B
road
,veryshort
––
–W
elld
evelope
d–
––
Cristainferior;po
orlyde
velope
d–
–
–Bridg
e-like
––
–Collu
m:A
bsen
t–
–
JAWAD 1055FISH
TABLE
7(Continue
d)
SolomonIsland
sArabian
–Persian
gulf
SouthAfrica
New
Caled
onia
Taiwan
–Dee
p,floorwithridg
es–
––
Cau
da:S
hallo
w,floorsm
ooth
––
–Dee
p,floorwithridg
es–
––
Ostium:S
hallo
w,floor
Withfine
lumps
––
–W
elld
evelope
d–
––
-–
–
Scarus
niger
161mm
L T161mm
L T202mm
L T157mm
L T158mm
L T206mm
L T
Oval
Welld
evelope
dW
ithseve
rallobe
sSh
ape:
Circu
lar
Collu
m:A
bsent
Posteriormargin:
Withonelobe
Lobe
dBridg
e-like
Broad
lypo
inted
Dorsalmargin:
Withinde
ntation
above
caud
aCristainferior:Poorly
dev
eloped
Antirostrum;finepointed
Coarse
lobe
dDee
p,floorwithridg
esW
ithco
arse
ridge
Ven
tralmargin;
fine
cren
ate
Ostium
andcauda:
Shallow
withsm
ooth
floor
Ostium
andcauda:
Smooth
withoneridge
Coarse
lobe
dAbsen
tW
elld
evelope
dPosteriormargin:
Curve
dVen
traldep
ression:S
hallow
Collu
m:P
oorlydev
eloped
Long
,lobe
d215mm
–Rostrum;sho
rt,roun
ded
220mm
–
Dee
p,floorwithridg
esSlightly
cren
ate
–Cau
da:S
hallo
w,floorsm
ooth
Ven
tralmargin:C
oarsely
lobed
–
Floorwithlumps
and
ridg
esat
bottom
Broad
lypo
inted
–Ostium:F
loorwithcentralridge
Posteriormargin;straigh
t–
215mm
L TBroad
lypo
inted
–286mm
L TRostrum:W
ithtw
osm
all
projections
–
Withbroad
fissure
Poorlyde
velope
d–
Posteriormargin:
Pointed
Antirostrum:S
hort,p
ointed
–
Long
,sha
llow
Floorwithlumps
–Dorsalde
pression:
Absen
tCau
da:
Floorwithridge
s–
Floorwithco
arse
lobe
san
dridg
es–
–Cau
daan
dostium:F
loorsm
ooth
––
Welld
evelope
d–
–Cristasupe
rior:Poorlyde
velope
d–
–
Scarus
rivulatus
217mm
L T261mm
L T291mm
L T225mm
L T
Shap
e:Domed
withlobe
sElliptic
Shap
e;oblong
Dorsalmargin:E
marginate
Dorsalmargin:
Withde
epfissure
Finecren
ate
Dorsalmargin:C
oarse
lobed
Posteriormargin:
Withshallow
fissure
Rostrum:N
arrow,
fine
pointed
Long
,broad
lylobe
d,lower
edge
coarsely
lobe
dRostrum:S
hort,b
oard,lower
edge
emarginate
Rostrum:B
road
lypointed
Ecisura:W
ith
deve
lope
dprojection
Widewithwideno
tch
Ecisura:N
arrow
with
narrow
notch
Antirostrum:S
hort,b
road
Floorwithlumps
and
ridg
esCau
daan
dostium:
Floorsm
ooth
1056 JAWADFISH
species of Merluccius (Merlucciidae) can be separated from the
American ones, based on the variation of the sulcus; therefore, the
variation in sulcus runs parallel to zoogeography and phylogeny
(Torres et al., 2000). However, the variation in sulcus is also the result
of an acoustic specialization and thus interspecific sulcus variation
may also result from ecomorphological adaptations (Ramcharitar et al.,
2004; Popper et al., 2005). The shape of the sulcus is important in
sciaenid otoliths (Nolf 1985). Apart from sulcus morphology, the cor-
relation between particular otolith features (e.g., the proportions of
rostrum and antirostrum) and biological functions (e.g., swimming abil-
ity, feeding, or other activities) has not yet been established (Popper
et al., 2005). Considering the variety of teleosts, there may be some
correlation between the length of the rostrum and the swimming abil-
ity (Nolf, 1985; Volpedo & Echeverría, 2003), but this correlation is
not significant in the discrimination of closely related species
(Reichenbacher et al., 2007). In this study, the general morphology of
the rostrum in the six species examined is broad, short and lobed and
such a consistency agrees with Reichenbacher et al. (2007).
Selective pressures can sometimes act on sagittae so that their
morphology meets specific auditory needs (Gauldie, 1988; Platt &
Popper, 1981; Popper & Coombs, 1982). Constraints in terms of phys-
ical packing of sagittae within the skull have also been cited in several
studies, especially those examining closely related species with large
sagittae (Arellano et al., 1995; Gaemers, 1984; Smith, 1992). Otolith
growth has long been shown to have a positive relationship to fish
age (Sabetian et al., 2015), but otoliths in very large fishes can be
much smaller than those in small fishes and vice versa (Campana,
2004). A limited comparison made by Friedland and Reddin (1994)
suggests that the shape of otolith is also influenced by genetics. Dif-
ferent morphologies in otoliths of a species obtained from different
localities have been observed. The otoliths of the six scarid species
were compared with those of similar size from other localities and this
comparison is given here (Table 7). Rivaton and Bourret (1999) pro-
vided an image of an otolith from a S. rivulatus of 470 mm length, no
comparable length group was available for the present study. The oto-
lith described by Rivaton and Bourret (1999) is oblong with dorsal
margins nearly smooth and ventral margins crenulated. The rostrum is
long and lobed, there is no collum, the cauda is deep with a smooth
floor and the crista superior extends over the whole length of the
ostium. The largest otolith of S. rivulatus obtained for the present
study is elliptic and there are more differences in several morphologi-
cal aspects. The description of the otolith by Rivaton and Bourret
(1999) complements our knowledge of the otoliths of S. niger.
The results of this study highlight a degree of intraspecific varia-
tion in the shape of the otolith for each species, such as the angle of
the rostrum (straight v. oblique), or the shape of the excisura
(Tables 1–6). This study was unable to highlight enough variability in
otolith shape to use as a key for individual species from the Solomon
Islands, but there is enough variability to distinguish small–juvenile
from large–adult individuals provided the species has been rigourously
identified upon capture. Further investigation is required, including a
comparative study of the shape and geometry of the sagittal otolith,
to add further taxonomic characters for the identification of these
species from the Solomon Islands.
ACKNOWLEDGEMENTS
My sincere thanks are due to those people who collected the fish
specimens and extracted their otolith. Also to the Auckland University
of Technology (AUT) for the SEM imaging of the otoliths and for edit-
ing them in Photoshop.
ORCID
Laith A. Jawad https://orcid.org/0000-0002-8294-2944
REFERENCES
Aguirre, H., & Lombarte, A. (1999). Ecomorphological comparisons of sagit-tae in Mullus barbatus and M. surmuletus. Journal of Fish Biology, 55,105–114.
Arellano, R. V., Hamerlynck, O., Vincx, M., Mees, J., Hostens, K., &Gijselinck, W. (1995). Changes in ratio of the sulcus acusticus area tothe sagitta area of Pomatoschistus minutus and P. lozanoi (Pisces, Gobii-dae). Marine Biology, 122, 355–360.
Assis, C. A. (2003). The lagenar otoliths of teleosts: Their morphology andits application in species identification, phylogeny and systematics.Journal of Fish Biology, 62, 1268–1295.
Baremore, I. E., & Bethea, D. M. (2010). A guide to otoliths from fishes of theGulf of Mexico. NOAA technical memorandum NMFS-SEFSC-599.Washington, DC: U.S. Department of Commerce. Retrieved fromwww.repository.library.noaa.gov/view/noaa/4010
Bellwood, D. R. (1994). Phylogenetic study of the parrotfishes family Scari-dae (Pisces: Labroidei), with a revision of genera. Records of theAustralian Museum, Supplement, 20, 1–86.
Campana, S. E. (2004). Photographic atlas of fish otoliths of the northwestatlantic Ocean Canadian Special publication of fisheries and aquatic sci-ences (Vol. 133). Ottawa, ON: NRC Research Press. Retrieved fromwww.dfo-mpo.gc.ca/Library/278479.pdf
Chaine, J., & Duvergier, J. (1934). Recherches sur les otoliths des poisons.Etude descriptive et comparative de la sagitta des teléostéens. Actesde la Société Linnéenne de Bordeaux, 86, 5–254.
Cruz, A., & Lombarte, A. (2004). Otolith size and its relationship with col-our patterns and sound production. Journal of Fish Biology, 65,1512–1525.
Friedland, K. D., & Reddin, D. G. (1994). Use of otolith morphology in stockdiscriminations of Atlantic salmon (Salmo salar). Canadian Journal ofFisheries and Aquatic Science, 51, 91–98.
Frizell, D. L., & Dante, J. H. (1965). Otoliths of some early Cenozoic fishesof the Gulf coast. Journal of Palaeontology, 39, 686–718.
Gaemers, P. A. M. (1984). Taxonomic position of the Cichlidae (Pisces, Per-ciformes) as demonstrated by the morphology of their otoliths. Nether-land Journal of Zoology, 34, 566–595.
Gauldie, R. W. (1988). Function, form and time-keeping properties of fishotoliths. Comparative Biochemistry and Physiology A, 91, 395–402.
Gauldie, R. W., & Nelson, D. G. A. (1990). Otolith growth in fishes. Compar-ative Biochemistry and Physiology A, 97, 119–135.
Ibsch, M., Anken, R. H., & Rahmann, H. (2004). Calcium gradients in thefish inner ear sensory epithelium and otolithic membrane visualized byenergy filtering transmission electron microscopy (EFTEM). Advances inSpace Research, 33, 1395–1400.
Jawad, L. A. (2008). Comparative morphology of the otolith of the triple-fins (family: Tripterygiidae). Journal of Natural History, 41, 901–924.
Jawad, L. A., Al-Jufaili, S. M., & Al-Shuhaily, S. S. (2007). Scale morphologyof greater lizardfish Saurida tumbil (Bloch, 1795) (Pisces: Synodonti-dae). Journal of Fish Biology, 70, 1185–1212.
Jawad, L. A., Hoedemakers, K., Ibáñez, A. L., Ahmed, Y. A.,El-Regal, M. A. A., & Mehanna, S. F. (2017). Morphology study of theotoliths of the parrotfish, Chlorurus sordidus (Forsskål, 1775) and Hip-poscarus harid (Forsskål, 1775) from the Red Sea coast of Egypt (family:Scaridae). Journal of the Marine Biological Association of the United King-dom, 98, 1–10. https://doi.org/10.1017/S0025315416002034
Klingenberg, C. P., & Ekau, W. (1996). A combined morphometric and phy-logenetic analysis of an ecomorphological trend: Pelagization in
L'Abée-Lund, J. H. (1988). Otolith shape discriminates between juvenileAtlantic salmon, Salmo salar L. and brown trout, Salmo trutta L. Journalof Fish Biology, 33, 899–903.
L'Abée-Lund, J. H., & Jensen, A. J. (1993). Otoliths as natural tags in thesystematic of salmonids. Environmental Biology of Fishes, 36, 389–393.
Ladich, F., & Popper, A. N. (2001). Comparison of the inner ear ultrastruc-ture between teleost fishes using different channels for communica-tion. Hearing Research, 154, 62–72.
Lin, C. H., & Chang, C. W. (2012). Otolith atlas of Taiwan fishes NMBBAatlas series (Vol. 12). Checheng, Taiwan: National Museum of MarineBiology and Aquarium.
Lombarte, A., & Fortuño, J. M. (1993). Differences in morphological fea-tures of the sacculus of the inner ear of two hakes (Merluccius capen-sis and M. paradoxus, Gadiformes) inhabits from different depth ofsea. Journal of Morphology, 214(1), 97–107.
Lombarte, A., Torres, G. J., & Morales-Nin, B. (2003). Specific Merlucciusotolith growth patterns related to phylogenetics and environmental.Journal of the Marine Biological Association of United Kingdom, 83,27781.
Martinez-Andrade, F. (2003). A comparison of life histories and ecologicalaspects among snappers (Pisces: Lutjanidae). (PhD thesis). Lousiana StateUniversity, Baton Rouge.
Nelson, J. S., Grande, T. C., & Wilson, M. V. H. (2016). Fishes of the world.Hoboken, NJ: Wiley & Sons Inc.
Nolf, D. (1985). Handbook of Paleoichthyology. Volume IO: Otolithi piscium.Stuttgart, Germany: Fischer Verlag.
Nolf, D. (2013). The diversity of fish otoliths, past and present. Brussels, Bel-gium: Royal Belgian Institute of Natural Sciences.
Pannella, G. (1980). Growth pattern of fish sagittae. In D. C. Rhoad &R. A. Lutz (Eds.), Skeletal growth of aquatic organisms: Biological recordsof environmental change, topics in geobiology (Vol. 1, pp. 519–560).New York, NY: Plenum Press.
Parenti, P., & Randall, J. E. (2011). Checklist of the species of the familiesLabridae and Scaridae: An update. Smithiana Bulletin, 13, 29–44.
Paxton, J. R. (2000). Fish otoliths: Do sizes correlate with taxonomic group,habitat and/or luminescence? Philosophical Transaction of the RoyalSociety A, 355, 1299–1303.
Platt, C., & Popper, A. N. (1981). Fine structure and function of the ear. InW. N. Tavolga, A. N. Popper, & R. R. Fay (Eds.), Hearing and sound com-munication in fish (pp. 3–38). New York, NY: Springer Verlag.
Ponton, D. (2006). Is geometric morphometrics efficient for comparingotolith shape of different fish species? Journal of Morphology, 267,750–757.
Popper, A. N., & Coombs, S. (1982). The morphology and evolution of theear in Actinopterygian fishes. American Zoologist, 22, 311–328.
Popper, A. N., & Hoxter, B. (1981). The fine structure of the sacculus andlagena of a teleost fish. Hearing Research, 5, 245–263.
Popper, A. N., Ramcharitar, J., & Campana, S. E. (2005). Why otoliths?Insights from inner ear physiology and fisheries biology. Marine andFreshwater Research, 56, 497–504.
Ramcharitar, J. U., Deng, X., Ketten, D., & Popper, A. N. (2004). Form andfunction in the unique inner ear of a teleost: The silver perch (Bairdiellachrysoura). Journal of Comparative Neurology, 475, 531–539.
Randall, J. E. (2005). Reef and shore fishes of the South Pacific: New Caledo-nia to Tahiti and the Pitcairn Islands. Honolulu, HI: University of HawaiiPress.
Reichenbacher, B., Sienknecht, U., K¨uchenhoff, H., & Fenske, N. (2007).Combined otolith morphology and morphometry for assessing taxon-omy and diversity in fossil and extant killifish (Aphanius, Prolebias).Journal of Morphology, 268, 898–915.
Rivaton, J., & Bourret, P. (1999). Institut de recherche pour le développ-ment. In Les otolithes des poissons de l'IndoPacifique. Documents
Scientifiques et Techniques 112. Nouméa, New Caledonia. Retrievedfrom www.horizon.documentation.ird.fr/exl-doc/pleins_textes/divers09-06/010021515.pdf
Ruck, J. G. (1976). Studies on the development and osteology of someNew Zealand inshore fishes. (PhD thesis). Victoria University, Wellington.
Sabetian, A., Trip, E. L., Wheeler, P., Sands, L., Wakefield, S., Visconti, V., &Banda, F. (2015). Biological plasticity of non-native European perch(Perca fluviatilis) populations and the implications for management innorthern New Zealand. Journal of the Marine and Freshwater Research,49, 119–131.
Sadighzadeh, Z., Otero-Ferrer, J. L., Lombarte, A., Fatemi, M. R., &Tuset, V. M. (2014). An approach to unraveling the coexistence ofsnappers (Lutjanidae) using otolith morphology. Scientia Marina, 78,353–362.
Sadighzadeh, Z., Tuset, V. M., Valinassab, T., Dadpour, M. R., &Lombarte, A. (2012). Comparison of different otolith shape descriptorsand morphometrics for the identification of closely related species ofLutjanus spp. from the Persian Gulf. Marine Biology Research, 8,802–814.
Schulz-Mirbach, T., & Reichenbacher, B. (2006). Reconstruction of oligi-cene and neogene freshwater fish faunas-an actualistic study on cypri-niform otoliths. Acta Palaeontologica Polonica, 51, 283–304.
Smale, M. J., Watson, G., & Hecht, T. (1995). Otolith atlas of southern Afri-can marine fishes, Ichthyological Monographs 1. Grahamstown,South Africa: Smith Institute of Ichthyology.
Smith, M. K. (1992). Regional differences in otolith morphology of thedeep slope red snapper Etelis carbunculus. Canadian Journal of Fisheriesand Aquatic Science, 49, 795–804.
Teimori, A., Jawad, L. A. J., Al-Kharusi, L. H., Al-Mamry, J. M., &Reichenbacher, B. (2012). Late Pleistocene to Holocene diversificationand historical zoogeography of the Arabian killifish (Aphanius dispar)inferred from otolith morphology. Scientia Marina, 76, 637–645.
Torres, G. J., Lombarte, A., & Morales-Nin, B. (2000). Variability of the sul-cus acusticus in the sagittal otolith of the genus Merluccius(Merlucciidae). Fisheries Research, 46, 5–13.
Tuset, V. M., Lombarte, A., & Assis, C. A. (2008). Otolith atlas for the west-ern Mediterranean, north and central eastern Atlantic. Scientia Marina,72, 7198.
Tuset, V. M., Lombarte, A., Gonzalez, J. A., Pertusa, J. F., & Lorente, M. J.(2003a). Comparative morphology of the sagittal otolith in Serranusspp. Journal of Fish Biology, 63, 1491–1504.
Tuset, V. M., Lozano, I. J., Gonzalez, J. A., Pertusa, J. F., &Garcia-Diaz, M. M. (2003b). Shape indices to identify regional differ-ences in otolith morphology of comber, Serranus cabrilla. Journal ofApplied Ichthyology, 19, 88–93.
Tuset, V. M., Rosin, P. L., & Lombarte, A. (2006). Sagittal otolith shape usedin the identification of fishes of the genus Serranus. Fisheries Research,81, 316–325.
Volpedo, A., & Echevarría, D. D. (2003). Ecomorphological patterns of theSagitta in fish on the continental shelf off argentine. Fisheries Research,60, 551–560.
Weisler, M. I. (1993). The importance of fish otoliths in Pacific IslandArchaeofaunal analysis. New Zealand Journal of Archaeology, 15,131–159.
How to cite this article: Jawad LA. A comparative morpho-
logical investigation of otoliths of six parrotfish species (Scari-
dae) from the Solomon Islands. J Fish Biol. 2018;93: