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A new genus and species of pomacentrid sh (Perciformes)
from the Eocene of Bolca in northern Italy
ALEXANDRE F. BANNIKOV*, DAVID R. BELLWOOD**
(*Borisyak Paleontological Institute of the Russian Academy of Sciences, Moscow, Russia)
(**School of Marine and Tropical Biology and ARC Centre of Excellence for Coral Reef Studies, James Cook University,
Townsville, Australia)
A
A new genus and species of pomacentrid sh, Sorbinichromis francescoi, is described based on a single skeleton from the Eocene locality
of Bolca in northern Italy. Sorbinichromis gen. nov. is characterized by a unique combination of vertebral (10+15=25) and unpaired
n-ray counts (D XI, 12; A II, 10; C 4, I,8-7,I, 3), long pelvic ns, and the possession of two supraneurals.
Key words: Perciformes, Pomacentridae, new genus and species, Eocene, northern Italy, Bolca locality.
R
Sorbinichromis francescoi, nuovo genere e nuova specie della famiglia Pomacentridae, è descritto sulla base di un singolo scheletro
dell’Eocene di Bolca (Italia settentrionale). Sorbinichromis francescoi è caratterizzato dalla combinazione unica del numero di
vertebre (10+15=25) e di raggi delle pinne impari (D XI, 12; A II, 10; C 4, I, 8 – 7, I, 3), dalle lunghe pinne pelviche, e dalla
presenza di due supraneurali.
Parole chiave: Perciformes, Pomacentridae, nuovo genere e nuova specie, Eocene, Italia settentrionale, Bolca.
I
e Early Eocene sh fauna of the famous locality
Monte Bolca, to the north of Verona in northern Ita-
ly, is exceptionally rich in marine shes, especially the
acanthopterygians. Monte Bolca marks the rst fossil
record of many groups of shes found on modern cor-
al reefs (Bellwood, ). Some groups of Recent sh-
es (e.g., butteryshes and gobies) rst appeared on-
ly in the Oligocene or later in the Eocene; these were
represented in the Bolca assemblage by their extinct
ecological analogues (Bannikov, a, b). Except for
discoveries of fragmentary remains (pharyngeal jaws)
elsewhere, the Bolca sh fauna establishes the earliest
known record of the Pharyngognathi, or Labroidei.
Bellwood () reviewed the status of the Bolca phar-
yngognath shes and found that the genera Odonte-
obolca Krell (=Odonteus Agassiz), Gillidia Eastman and
Eolabroides Eastman appear to be non-pharyngognaths,
whereas Labrus ? valenciennesi Agassiz is an indetermi-
nate non-labroid pharyngognath. He stated (Bellwood,
) that the Bolca sh fauna contains at least six la-
broid taxa: two monotypic genera in the Labridae (Eo-
coris Bannikov et Sorbini and Phyllopharyngodon Bell-
wood), two monotypic genera in the Pomacentridae
(Palaeopomacentrus Bellwood et Sorbini and Lorenzich-
thys Bellwood), as well as a range of other labroids of
less certain anity, including the Tortonesidae (mono-
typic family established for the genus Tortonesia Sorb-
ini: Sorbini et al., ) and the genus Sorbinia Bell-
wood. e latter (with type species S. caudopunctata)
was described by Bellwood () based on a single
specimen with a somewhat damaged caudal n. Bell-
wood () indicated the caudal-n formula of the
holotype as +-?+. Subsequently seven more speci-
mens of S. caudopunctata were located in the collec-
tion of the Museo Civico di Storia Naturale di Verona
(MCSNV) in addition to a specimen in the private col-
lection of the family Cerato in Bolca (gured in Cer-
ato, ): MCSNV , MCSNV T/T, MC-
SNV T/T, MCSNV T, MCSNV IG/
IG, MCSNV IG/ IG, and MCS-
NV IG (the latter specimen was erroneously clas-
sied with the Mullidae: Blot, : ; see Bannikov,
). Most of these have a well-preserved caudal n,
and the caudal-n formula of S. caudopunctata should
be regarded as -, I+-+I, .
Following the review of the Bolca labroids (Bell-
wood, ), a new genus and species Quasicichla mu-
cistonaver was described and interpreted as closely relat-
ed to the family Cichlidae (Bannikov, b). Recently,
the third taxon attributable to the Labridae, Bellwood-
Studi e ricerche sui giacimenti terziari di Bolca, XV - Miscellanea paleontologica, 12, 2014: 7 - 14
8
ALEXANDRE F. BANNIKOV - DAVID R. BELLWOOD
ilabrus landinii Bannikov et Carnevale, was described
from the Monte Bolca locality, Pesciara cave site (Ban-
nikov and Carnevale, ).
Among the putative labroids held in the collection
of the MCSNV there are three specimens of the sh
gured by Caltran and Zorzin () and provisional-
ly placed in the labroid family Embiotocidae. is sh
was subsequently described as an incertae sedis percoid
Frippia labroiformis (Bannikov and Carnevale, ).
Revision of the morphology and systematic position
of Bradyurus (=Eolabroides) szajnochae (de Zigno, )
conrmed the conclusion of Bellwood () that this
taxon is non-pharyngognath; it is regarded as a percoid
of uncertain anity, although it may have haemulid re-
lationships (Bannikov and Zorzin, ).
Given the growing evidence that the Labroidei sen-
su lato is not monophyletic, Bannikov and Carnevale
() proposed that the labroid anities of Quasicich-
la, Sorbinia and Tortonesia should be reconsidered. e
Pomacentridae, although pharyngognath, are no long-
er regarded as labroids and recent molecular evidence
suggests a distant relationship with other labroids, al-
though the close relationship with the Cichlidae re-
mains (Near et al., )
Of the two pomacentrid species recorded from Bol-
ca, Palaeopomacentrus orphae was originally described
(Bellwood and Sorbini, ) based on two specimens
(MCSNV IG Holotype; BMNH P Paratype).
e specimens were relatively intact with traces of pig-
mentation, including a dorsal ocellus reminiscent of
those on Recent species in the family. e second poma-
centrid species Lorenzichthys olihan (Bellwood ) was
based on a single specimen, although the head area was
poorly preserved. A distorted specimen MCSNV T
(in the same slab with the percoid Acropoma lepidotum),
and somewhat damaged MCSNV IG may also be-
long to Palaeopomacentrus orphae.
Among the fossil shes housed in the collection of
the MCSNV there is a specimen which can be inter-
preted as belonging to a new genus and species of the
Pomacentridae. It is the third damselsh taxon known
from the sh fauna of Monte Bolca. is sh is de-
scribed below.
M
e specimen was examined using a stereomicro-
scope with attached camera lucida drawing arm. Some
details of the specimen examined were best seen when
the specimen was moistened with alcohol. e speci-
men was prepared by needle.
Interneural and interhaemal spaces are numbered
based on the vertebra whose neural or haemal spine
forms the anterior border of the space, with the rst
space being between the rst and second neural or hae-
mal spines (following Baldwin and Johnson, ; Ban-
nikov and Tyler, ; Tyler and Bannikov, ; etc.).
Abbreviations are as follows: Institutional: MCSNV
- Museo Civico di Storia Naturale di Verona; Anatomi-
cal: HL - head length; PU - preural vertebra; SL - stand-
ard length; U - ural vertebra.
S D
Order Perciformes
Family Pomacentridae Girard,
Genus Sorbinichromis gen. nov.
Diagnosis
Relatively deep-bodied sh with a short and deep cau-
dal peduncle. Head relatively small. Maximum body
depth strongly exceeds head length. Supraoccipital crest
deep and pointed. Lower jaw articulation under mid-
dle of orbit. Mouth relatively small. (+) verte-
Fig. - Sorbinichromis francescoi gen. et sp. nov., holotype, part and
counterpart: A - MCSNV IG, B - MCSNV IG; Low-
er Eocene of Bolca in northern Italy, Pesciara. Scale division: mm.
9
A new genus And species of pomAcentrid fish (perciformes) from the eocene of BolcA in northern itAly
brae. Supraneurals . Dorsal n long-based and con-
tinuous, with strong spines and soft rays. ere
is slight notch between spiny and soft portions of dor-
sal n. Anal n with supernumerary spines (nd very
strong) and rays. Postcleithrum strong. Pelvics in-
serted just behind pectorals, strongly elongated. Cau-
dal n truncated. Scales relatively large, thick at dorsal
and ventral contours of body. Lateral line follows dor-
sal contour of body and interrupted below anterior por-
tion of soft dorsal n.
Type Species
Sorbinichromis francescoi sp. nov., by monotypy and
designation herein.
Etymology
e genus is named in honour of the late Lorenzo Sorb-
ini, preeminent Italian paleoichthyologist and a dear
friend of both authors; and the genus Chromis; gen-
der masculine.
Composition
Type species only.
Sorbinichromis francescoi sp. nov.
Figures ,
Diagnosis
As for the genus.
Etymology
e species is named in honour of the son of Loren-
zo Sorbini, Francesco, who kindly made the photos of
Bolca shes for this and many other papers.
Holotype
MCSNV IG/, part and counterpart, com-
plete skeleton relatively poorly preserved and somewhat
damaged by fractures, mm SL.
Referred Specimens
None.
Type Locality And Horizon
Monte Bolca locality, Pesciara cave site; Early Eocene,
late Ypresian, middle Cuisian, SBZ , Alveolina dainel-
lii Zone (see Papazzoni and Trevisani, ).
Fig. - Sorbinichromis francescoi gen. et sp. nov., reconstruction of the skeleton based on holotype, scales omitted.
10
ALEXANDRE F. BANNIKOV - DAVID R. BELLWOOD
Description
e body is relatively deep, with a short and deep cau-
dal peduncle. e body depth is less than half the SL.
e caudal peduncle depth is about . of the body
depth. e head is relatively small; its length is about
. times in the body depth. e head length is con-
tained approximately . times in SL. e dorsal pro-
le of the body is almost equally as convex as the ven-
tral prole of the body.
Head. e head is slightly deeper than long. e or-
bit is moderate and placed relatively high on the head
depth. e horizontal diameter of the orbit is about -
HL. e snout is slightly shorter than the orbit
diameter. e mouth is relatively small and terminal.
e lower jaw articulation is situated below the middle
of the orbit. Faint remains of some infraorbital bones
bearing suborbital branch of the lateral line sensory ca-
nal are recognizable in the holotypic counterpart MC-
SNV IG . e neurocranium is relatively deep,
with the supraoccipital crest strongly developed, trian-
gular and pointed. e ethmoid region is short. e
parasphenoid is strongly fragmented; it seems to pass at
the lower border of the orbit. e upper jaw bones are
incompletely preserved, with the premaxilla best seen
in MCSNV IG , whereas the maxilla in MCS-
NV IG . e premaxilla has a moderate ascend-
ing process divided from an articular process. Few pre-
maxillary teeth are preserved, these are small, slender
and unicuspid. No supramaxilla is evident. e lower
jaw is badly damaged; its length is about - HL.
e dentary has oblique symphysis. e dentition of
the dentary is equal to that of the upper jaw; the small
sharp teeth being dark at their tips. e hyomandibu-
lar shaft is slightly inclined from the vertical line. De-
tails of the structure of the suspensorium and pterygoi-
ds are not clear. e thin endopterygoid forms the oor
of the orbit. e opercular region is rather broad, with
individual bones being indistinct. None of the branchi-
al and hyoid bones are recognizable, except for the faint
remains of what seems to be the lower pharyngeal jaw
in MCSNV IG . No pharyngeal teeth are evident.
Axial skeleton. ere are most probably vertebrae,
including the urostyle: ten abdominal and fteen cau-
dal (the anteriormost vertebrae are hardly recogniza-
ble). e axis of the vertebral column is sigmoid and
elevated anteriorly. e vertebral centra are mostly rec-
tangular in lateral view, they bear poorly developed
longitudinal lateral ridge on each side. e length of
the caudal portion of the vertebral column is . times
greater than the length of the abdominal portion of the
vertebral column. e vertebral spines are moderately
long and robust, straight or slightly curved. e neu-
ral spines of most of the anterior abdominal vertebrae
are more strongly expanded than those of the succeed-
ing vertebrae. e haemal spines of most of the caudal
vertebrae are longer and more strongly inclined to the
axis of the backbone than the opposite neural spines.
e parapophyses are poorly recognizable in the poste-
riormost abdominal vertebrae. e pleural ribs are long
and rather slender, these rather strongly inclined pos-
teroventrally. None of epineurals are recognizable.
Pectoral n and girdle. Most of the pectoral girdle
bones are either not preserved or scarcely recogniza-
ble; however, the obliquely directed supracleithrum is
distinct, being strong and elongate. e ventral post-
cleithrum is also clearly visible; it is long and broad,
extending posteroventrally and terminating behind
the pelvic-n base. e pectoral n is not preserved;
its base can be tentatively reconstructed somewhere be-
low the fth vertebra in the middle of the distance be-
tween the vertebral column and the ventral prole of
the body, probably just anterior to the pelvic n.
Pelvic n and girdle. e pelvic bones are robust.
e pelvic ns contain a spine and probably ve soft,
branched rays. e pelvic n is long; its longest ray
reaches well behind the anal-n origin. e pelvic-n
spine is long but much shorter than the longest pelvic-
n soft rays; it seems to be no shorter than the longest
dorsal-n spine.
Supraneurals and dorsal n. ere are two relative-
ly slender supraneurals with minute apical projections.
e supraneurals seem to be accommodated between
the supraoccipital and the neural spine of the rst verte-
bra, i.e., in preneural space. e dorsal n is long-based
and continuous, with a slight notch between its spiny
and soft portions; it originates over the second verte-
bra and terminates over the -th vertebra. ere are
dorsal-n spines and soft segmented and branched
rays. e dorsal-n spines are relatively strong. e sec-
ond spine is the longest, subsequent spines gradual-
ly decrease in length posteriorly in the series. e sec-
ond spine is . times longer than the rst spine and
. times longer than the last spine. e rst dorsal-n
spine is supernumerary on the rst dorsal-n pterygio-
phore. e longest soft ray of the dorsal n (situated in
the middle of the soft portion) is . times longer than
the last dorsal-n spine. e length of the base of the
soft portion of the dorsal n is . times shorter than
the base length of the spiny portion of the dorsal n.
ere are a total of dorsal-n pterygiophores. e
rst pterygiophore is large and sturdy, expanded an-
teroposteriorly, and bears a longitudinal strengthening
ridge and procumbent distal process; the succeeding
pterygiophores of spines are also broad, whereas those
11
A new genus And species of pomAcentrid fish (perciformes) from the eocene of BolcA in northern itAly
of soft rays become narrower and decrease in length
posteriorly in the series. e interneural spaces below
the dorsal n have the ventral shafts of one (most of the
n) or two (four spaces posteriorly) pterygiophores pre-
sent. ere are no vacant interneural spaces.
Anal n. e anal n is relatively short at the base;
it originates under the fth caudal vertebra and termi-
nates under the end of the dorsal n. ere are two
spines and ten soft, segmented and branched rays in
the anal n. e rst anal-n spine is weak, whereas the
second spine is very strong, being slightly longer than
the longest (second) dorsal-n spine. Both two anal-n
spines are supernumerary. e longest (anterior) anal-
n soft rays are longer than the longest dorsal-n soft
rays. e rst anal-n pterygiophore is very long and
sturdy, wedge-like in shape; it is inclined at an angle
š to the body axis. e succeeding anal-n pterygio-
phores are much shorter and slender; these decrease in
length posteriorly in the series.
Caudal n and skeleton. e caudal skeleton is badly
damaged by two fractures in the matrix. It is clear that
terminal centrum is composed of the fusion of PU, U
and U; however, conditions of the hypurals and pa-
rhypural are less easily recognizable. e neural spine of
PU seems to be a short crest. ere are at least two epu-
rals, the rst of which is curved and longest. e caudal
n is truncated, moderately long but deep. We recon-
struct principal rays in the caudal n (I,-,I); there
are four procurrent rays above and three rays below.
Squamation. Large and thick scales cover the entire
body and the head. Some scales exhibit basal radii and
concentric striations; however, ctenii were not observed
at all (but there are no scales in the material with a well-
preserved apical eld). Scales are especially thick at the
dorsal and ventral contours of the body. Posterior por-
tion of the lateral line courses relatively high, it follows
the dorsal prole of the body and is interrupted below
the anterior third of the soft portion of the dorsal n.
No pigmentation is traceable.
Measurements. As percentage of SL of holotype:
head length = ; maximum body depth = ; caudal
peduncle depth = ; snout length = ; orbit diameter
= ; lower jaw length = ; distance between tip of the
snout and rst dorsal-n spine = ; distance between
tip of the snout and rst dorsal-n soft ray = ; dis-
tance between tip of the snout and anal n = .; dis-
tance between pelvic and anal ns = ; dorsal-n base
length = .; spinous dorsal-n base length = ; anal-
n base length = ; length of the rst dorsal-n spine =
; length of the longest dorsal-n spine = ; length of
the second anal-n spine = ; length of the pelvic n =
; length of the longest caudal-n ray = .
D
Sorbinichromis francescoi is the third pomacentrid
fossil to be recorded from Bolca. is species, along with
Palaeopomacentrus orphae and Lorenzichthys olihan, also
from Bolca, represent the entire pomacentrid fossil re-
cord to date (based on intact material) from the Paleo-
gene. Other pomacentrid material is restricted to Izuus
and Chromis from the Miocene of Japan and Algeria, re-
spectively (Bellwood and Sorbini, ). e skull of a
Pomacentridae indet. was described from the Miocene
of Italy (Carnevale and Landini, ). e imprint
of an incomplete skeleton was described by Prokof-
iev () from the Upper Oligocene or Lower Mio-
cene of Azerbaijan (erroneously indicated by Prokoev
as Dagestan, Russia), designated as Pomacentridae gen.
et sp. indet. However, the specimen lacks the important
pomacentrid synapomorphy, possession of two super-
numerary anal-n spines (Bellwood and Sorbini, ),
and cannot be placed in the family (see Bannikov, ).
Recognition of Recent families in the fossil record
can be fraught with diculties, often because of a lack
of suitable morphological synapomorphies (Bannik-
ov, b; Bellwood and Sorbini, ). Fortunately,
the Pomacentridae has at least one good synapomorphy
that is usually well preserved: two supernumerary anal-
n spines (Bellwood and Sorbini, ). While not
unique (also found in the Apogonidae), in combination
with other features it enables specimens to be clearly
separated from other families, including the closely-re-
lated Cichlidae (which usually has anal spines) (dis-
cussed in Bellwood and Sorbini, ). e general
morphology, putative pharyngeal jaw, and two super-
numerary anal spines strongly support the placement
of Sorbinichromis in the Pomacentridae.
Other features and meristic counts of Sorbin-
ichromis preclude placement in any Recent genus. Sorb-
inichromis has fewer vertebrae and fewer spines and rays
than most extant taxa, although it does overlap in mer-
istics with Chrysiptera (D XI-XIV, -; A II, -; Ta-
ble ). It may, however, be separated from most extant
pomacentrids in the possession of vertebrae (most
extant forms have vs. in Sorbinichromis) and two
supraneurals (a condition previously recorded only in
Premnas and Acanthochromis (Fitzpatrick, ; Table
). Furthermore, in Sorbinichromis the supraneurals are
unusual in that they appear to lie in front of the neural
spines, whereas in the Recent taxa with two supraneu-
rals the posteriormost supraneural inserts between the
rst two neural spines (Fitzpatrick, ). Like many
other groups (Siganidae, Acanthuridae, Ephippidae)
Sorbinichromis may be reliably placed within an extant
12
ALEXANDRE F. BANNIKOV - DAVID R. BELLWOOD
family but it represents a distinct new genus with no
Recent representatives.
e presence of a third pomacentrid genus in Bol-
ca (at about - Ma) is consistent with molecular ev-
idence of early diversication of the Pomacentridae in
the Eocene (Cooper and Westneat, ; Cowman and
Bellwood, , ). Furthermore, it adds weight to
the suggestion that the biogeographic location for the
origins of the major pomacentrid lineages was, as in
many other taxa (Renema et al., ), in the Tethys
rather than the Indo-Australian Archipelago as suggest-
ed by some reconstructions from molecular phyloge-
nies (Cowman and Bellwood, ).
e general body shape and ns of Sorbinichromis
are similar to Recent species that hang, often in schools,
above the reefs such as its namesake Chromis and oth-
er genera including Neopomacentrus, Amblyglyphidio-
don, and Chrysiptera. In Sorbinichromis, the small head,
small teeth and apparent absence of robust teeth in the
pharyngeal (in contrast to Palaeopomacentrus) are all
consistent with feeding on small, possibly planktonic
material, while the deep body and deep caudal pedun-
cle are indicative of relatively slow swimming speeds.
More specically, it was likely to be fast turning and fast
accelerating, but with relatively slow cruising speeds.
e peduncle depth is strongly correlated with caudal
n aspect ratio in surgeonshes (Bellwood et al., ),
with aspect ratios being a widely recognized indicator
of swimming ability (high aspect ratio and narrow pe-
duncle being indicative of high cruising speeds vs. low
aspect ratio and deep peduncles being indicative of rap-
id acceleration but limited sustained swimming abili-
ties) cf. Bellwood and Wainwright (). Taken in
concert, the morphology of Sorbinichromis strongly re-
sembles modern Chrysiptera, Chromis and Amblyglyphi-
dodon suggesting that it was a planktivore or omnivore
living in close proximity to a complex hard substratum.
is trophic categorization of Sorbinichromis is con-
sistent with the suggestion that the shes of Monte Bolca
mark the initial record of an expansion in high-precision
diurnal feeding on benthic hardgrounds or coral reefs
(Goatley et al., ). is may have also included ben-
thic planktivores such as Sorbinichromis. Interestingly,
while some sh groups in the Eocene exhibit morphol-
ogies that are indicative of extensive trophic divergence
(e.g. the Acanthuridae, Bellwood et al., ) the poma-
centrids show relatively little divergence, a pattern that is
reected in extant forms (Cooper and Westneat, ).
e three Bolca pomacentrid genera, although taxonom-
ically dierent from their extant counterparts, share with
modern forms a limited expression in terms of morpho-
logical and trophic specialization.
Overall, the description of a third pomacentrid ge-
nus, Sorbinichromis, highlights the importance of Bol-
ca in the early diversication of modern sh groups.
e Pomacentridae is now equal with the Labridae with
three described genera. ey are thus among the most
diverse families within this Lagerstätte, certainly among
the pharyngognath representatives.
A
We are very grateful to Dr. Giuseppe Minciotti, Di-
rector of the MCSNV for providing funding for the
travel of AFB in to Verona to engage in this and
other studies. We also thank Dr. Roberto Zorzin and
Dr. Anna Vaccari at the MCSNV for their continuing
help in facilitating our research on the shes of Mon-
te Bolca; Dr. Leonardo Latella for editorial process-
ing of the manuscript; Christopher Goatley for assis-
tance with extant specimens, and Mrs. Bruna Burato
for her aid with the literature on Bolca shes. e re-
search of AFB was supported by the Russian Founda-
tion for Basic Research, project no. --. Prof.
David Bellwood was supported by the Australian Re-
search Council. Dr. Chiara Sorbini of the University of
Pisa translated the Abstract into Italian and Mr. Franc-
esco Sorbini (Verona) kindly made the photographs.
L C
ALLEN G.R. . Damselshes of the World. Aquar-
ium systems, Melle: pp.
ALLEN G.R., ERDMANN M.V. . Reef shes
of the East Indies. University of Hawai’i Press, Honolu-
lu: pp.
BALDWIN C.C., JOHNSON G.D., . Phylog-
eny of the Epinephelidae (Teleostei: Serranidae). Bulle-
tin of Marine Science (): -.
BANNIKOV A.F., a. Eocottidae, a new family
of perciform shes (Teleostei) from the Eocene of north-
ern Italy (Bolca). Miscellanea Paleontologica n. . Studi e
Ricerche sui Giacimenti Terziari di Bolca, : -.
BANNIKOV A.F., b. Fishes from the Eocene
of Bolca, northern Italy, previously classied with the
Chaetodontidae (Perciformes). Miscellanea Paleontolog-
ica n. . Studi e Ricerche sui Giacimenti Terziari di Bol-
ca, : -.
BANNIKOV A.F., . Two new genera for long
known percoid shes (Perciformes) from the Eocene
of Bolca, Italy. Miscellanea Paleontologica n. . Studi e
Ricerche sui Giacimenti Terziari di Bolca, : -.
13
A new genus And species of pomAcentrid fish (perciformes) from the eocene of BolcA in northern itAly
BANNIKOV A.F., . Fossil Vertebrates of Russia
and Adjacent Countries. Fossil Acanthopterygians Fishes
(Teleostei, Acanthopterygii). GEOS, Moscow: LXI+
pp. [In Russian].
BANNIKOV A.F., CARNEVALE G., . Bell-
woodilabrus landinii n. gen., n. sp., a new genus and
species of labrid sh (Teleostei, Perciformes) from the
Eocene of Monte Bolca. Geodiversitas, (): -.
BANNIKOV A.F., CARNEVALE G., . Frippia
labroiformis n. gen. n. sp., a new perciform sh from
the Eocene of Pesciara di Bolca, Italy. Bollettino della
Società Paleontologica Italiana, (): -.
BANNIKOV A.F., TYLER J.C., . Phylogenetic
revision of the sh families Luvaridae and †Kushluki-
idae (Acanthuroidei), with a new genus and two new
species of Eocene luvarids. Smithsonian Contributions
to Paleobiology, : -.
BANNIKOV A.F., ZORZIN R., (). Re-
description of the percoid sh Bradyurus (=Eolabroides)
szajnochae (de Zigno) (Perciformes, Percoidei) from the
Eocene of Bolca in northern Italy. Miscellanea paleon-
tologica n. . Studi e Ricerche sui Giacimenti Terziari di
Bolca, : -.
BELLWOOD D.R., (). A new Eocene fos-
sil sh, Sorbinia caudopunctata gen. et sp. nov., from
Monte Bolca, Italy. Bollettino del Museo Civico di Storia
Naturale di Verona, : -.
BELLWOOD D.R., . e Eocene shes of
Monte Bolca: the earliest coral reef sh assemblage.
Coral Reefs, : -.
BELLWOOD D.R., . Fossil pharyngognath
shes from Monte Bolca, Italy, with a description of a
new pomacentrid genus and species. Miscellanea Pale-
ontologica. Studi e Ricerche sui Giacimenti Terziari di
Bolca, : -.
BELLWOOD D.R., GOATLEY C.H.R., BRANDL
S.J., BELLWOOD O., . Fifty million years of her-
bivory on coral reefs: fossils, sh and functional inno-
vations. Proceedings of the Royal Society B, Biological Sci-
ences, : .
BELLWOOD D.R., SORBINI L., . A review
of the fossil record of the Pomacentridae (Pisces: La-
broidei) with a description of a new genus and species
from the Eocene of Monte Bolca, Italy. Zoological Jour-
nal of the Linnean Society, : -.
BELLWOOD D.R., WAINWRIGHT P.C., .
Locomotion in labrid shes: implications for habitat
use and cross-shelf biogeography on the Great Barrier
Reef. Coral Reefs, : -.
BLOT J., . La faune ichthyologique des gise-
ments du Monte Bolca (Province de Vérone, Italie).
Catalogue systématique présentant l’état actuel des re-
cherches concernant cette faune. Bulletin du Muséum
national d’Histoire naturelle (Paris), sér. , section C,
(): -.
CALTRAN T., ZORZIN R., . Bolca ed il suo
territorio: storia, tradizione, cultura e scienza. Golden
Time Communication, S. Maria di Zevio (VR). pp.
CARNEVALE G., LANDINI W., . A fossil
damselsh (Pisces, Pomacentridae) from the Late Mi-
ocene of Central Italy. Biological and biogeographical
considerations. Palaeontographia Italica, : -.
CERATO M., . Cerato. I pescatori del tempo.
Graca Alpone srl, San Giovanni Ilarione (VR). pp.
COOPER W.J., WESTNEAT M.W., . Form
and function of damselsh skulls: rapid and repeat-
ed evolution into a limited number of trophic niches.
BMC Evolutionary Biology, : .
COWMAN P.F., BELLWOOD D.R., . Coral
reefs as drivers of cladogenesis: expanding coral reefs,
cryptic extinction events, and the development of bi-
odiversity hotspots. Journal of Evolutionary Biology, :
-.
COWMAN P.F., BELLWOOD D.R., . e
historical biogeography of coral reef shes: global pat-
terns of origination and dispersal. Journal of Biogeogra-
phy, : -.
FITZPATRICK S., . Pomacentrid intrafamilial
relationships: a cladistic approach. Unpublished Hon-
ours esis, James Cook University, Townsville, Aus-
tralia: pp.
GOATLEY C.H.R., BELLWOOD D.R., BELL-
WOOD O., . Fishes on coral reefs: changing roles
over the past million years. Paleobiology, : -.
LEIS J.M., RENNIS D.S., . e larvae of In-
do-Pacic coral reef shes. New South Wales Universi-
ty Press, Sydney, University of Hawai’i Press, Honolu-
lu: pp.
NEAR T.J., DORNBURG A., EYTAN R.I., KE-
CK B.P., SMITH W.L., KUHN K.L., MOORE J.A.,
PRICE S.A., BURBRINK F.T., FRIEDMAN M.,
WAINWRIGHT P.C., . Phylogeny and tempo of
diversication in the superradiation of spiny-rayed sh-
es. Proceedings on the National Academy of Sciences of the
United States of America, : -.
PAPAZZONI C.A., TREVISANI E., . Facies
analysis, palaeoenvironmental reconstruction, and bi-
ostratigraphy of the “Pesciara di Bolca” (Verona, north-
ern Italy): An early Eocene Fossil-Lagerstätte. Palaeoge-
ography, Palaeoclimatology, Palaeoecology, :-.
PROKOFIEV A.M., . New and little known
perciform shes from the Upper Oligocene – Lower Mi-
ocene boundary deposits of the Caucasus (Osteichthyes,
Perciformes). Zoosystematica Rossica, (): -.
14
ALEXANDRE F. BANNIKOV - DAVID R. BELLWOOD
RENEMA W., BELLWOOD D.R., BRAGA J.C.,
BROMFIELD K., HALL R., JOHNSON K.G.,
LUNT P., MEYER C.P., MCMONAGLE L.B., MOR-
LEY R.J., O’DEA A., TODD J.A., WESSENLINGH
F.P., WILSON M.E.J., PANDOLFI J.M., . Hop-
ping hotspots: global shifts in marine biodiversity. Sci-
ence, : -.
SORBINI L., BOSCAINI E., BANNIKOV A.F.,
(). On the morphology and systematics of the
Eocene sh genus Tortonesia Sorbini from Bolca. Mis-
cellanea Paleontologica. Studi e Ricerche sui Giacimenti
Terziari di Bolca, : -.
TYLER J.C., BANNIKOV A.F., . Relation-
ships of the fossil and Recent genera of rabbitshes
(Acanthuroidei: Siganidae). Smithsonian Contributions
to Paleobiology, : -.
A
ALEXANDRE F. BANNIKOV
Borisyak Paleontological Institute of the Russian Acad-
emy of Sciences
Profsoyuznaya
Moscow
Russia
e-mail: aban@paleo.ru
DAVID R. BELLWOOD
School of Marine and Tropical Biology and ARC Cen-
tre of Excellence for Coral Reef Studies
James Cook University
Townsville, Queensland
Australia
e-mail: david.bellwood@jcu.edu.au
Table 1. Meristic values of Recent Pomacentridae.
Values drawn from Leis and Rennis (), Allen (),
Fitzpatrick () and Allen and Erdman (). Num-
bers refer to the number of species examined, numbers
in parentheses refers to the number of species for super-
aneurals in Fitzpatrick ().
Genus Dorsal n Anal Fin Vertebrae Supraneurals n
Abudefduf XIII, 11-17 II, 10-15 3 18 (4)
Acanthochromis XVII, 14-16 II, 14-16 2 1 (1)
Altrichthys XIV, 12-14 II, 14-15 2
Amblyglyphidodon XI-XIV, 10-16 II, 11-15 3 10 (2)
Amblypomacentrus XIII, 10-12 II, 11-13 2
Amphiprion VIII-XI, 13-20 II, 11-15 11 + 15 = 26 3 27 (3)
Azurina XII-XIII, 10-12 II, 11-13 2
Cheiloprion XIII, 13-14 II, 13-14 1
Chromis XII-XV, 9-15 II, 9-16 11 + 15 = 26 3 88 (6)
Chrysiptera XI-XIV, 9-15 II, 10-16 3 32 (3)
Dascyllus XII, 11-16 II, 11-16 3 9 (3)
Dischistodus XIII, 13-16 II, 13-16 3 6 (2)
Hemiglyphidodon XIII, 13-15 II, 13-15 1
Hypsypops XII, 16-17 II, 13-14 1
Lepidozygus XII, 14-15 II, 15-16 11 + 15 = 26 3 1 (1)
Mecaenichthys XIII, 16-17 II, 14 1
Microspathodon XII, 14-17 II, 12-14 3
Neoglyphidodon XIII-XIV, 12-16 II, 12-15 3 9 (1)
Neopomacentrus XIII, 10-13 II, 10-14 3 15 (2)
Nexilosus XIII, 17-19 II, 13-14 1
Parma XIII-XIV, 16-21 II, 13-16 10
Plectroglyphidodon XII-XIII, 14-20 II, 11-18 3 9 (1)
Pomacentrus XIII-XV, 12-16 II, 12-17 11 + 15 = 26 3 62
Pomachromis XIV, 12-14 II, 12-13 4
Premnas IX-X, 16-19 II, 13-15 2 1 (1)
Pristotis XIII, 12-13 II, 12-14 11 + 15 = 26 2
Similparma XIII, 17-18 II, 14 1
Stegastes XII-XIV, 12-17 II, 11-15 3 33 (1)
Teixeirichthys XIII, 11-13 II, 13 1
