Giuseppe Marramà

An Eocene anchovy from Monte Bolca, Italy: The earliest known record for the family Engraulidae

Engraulids, also known as anchovies, are a distinctive group of clupeoid fishes characterized by a series of derived morphological features of the snout and infraorbital bones, suspensorium and branchial arches. Although anchovies are very abundant today, they are scarcely represented in the fossil record. A new genus and species of anchovy, † Eoengraulis fasoloi gen. et sp. nov., is described from the Eocene (late Ypresian, c . 50 Ma) locality of Monte Bolca, Italy. It is based on a single well-preserved articulated skeleton that exhibits a unique combination of characters that supports its recognition as a new genus of the family Engraulidae, including: nine branchiostegal rays; 40 preural vertebrae and 17 pairs of pleural ribs; pleural ribs – preural vertebrae ratio 0.42; seven supraneurals; dorsal-fin origin at about mid-length of the body; about 16 dorsal-fin rays; anal-fin origin slightly behind the base of the last dorsal-fin ray; 19 anal-fin rays; seven pelvic-fin rays; and small needle-like pre-pelvic scutes. The morphological structure of the single available specimen suggests that † Eoengraulis fasoloi is the sister taxon of all other engrauline taxa. † Eoengraulis fasoloi is the oldest member of the family Engraulidae known to date. This taxon suggests that the earliest phases of diversification of engrauline anchovies probably occurred in the coastal palaeobiotopes of the western Tethys during Eocene time.

The Eocene sardine †Bolcaichthys catopygopterus (Woodward, 1901) from Monte Bolca, Italy: osteology, taxonomy, and paleobiology

ABSTRACT Clupeid fishes are abundant in the Eocene fossiliferous limestone of Monte Bolca, representing by far the most common group from this celebrated locality. However, despite of their abundance, the clupeid fishes from Monte Bolca have seldom been investigated. An analysis of about 300 well-preserved clupeid specimens from Monte Bolca housed in several institutions clearly indicates that more than 95% of the available material belongs to different developmental stages of a single taxon, traditionally referred to as †Clupea catopygoptera Woodward. †Bolcaichthys, a new genus of clupeid fish, erected to contain †Clupea catopygoptera Woodward from the Eocene of Monte Bolca, Italy, is described and compared with extant and fossil genera of the family Clupeidae. This new genus shows a unique combination of characters (head length approximately one third to one quarter standard length; skull roof with 10–14 frontoparietal striae; mouth terminal; two supramaxillae; edentulous jaws and palate; complete series of about 20–22 abdominal keeled scutes with ascending arms; no dorsal scutes; five or six branchiostegal rays; eight supraneurals; 40–42 vertebrae and 20–22 pleural ribs; three epurals) supporting its recognition as a new genus of the family Clupeidae. Paleobiological considerations suggest that the presence of a very large number of schooling clupeids belonging to the genus †Bolcaichthys at all stages of development (larval, juvenile, and adult) support the hypothesis that the sediments were deposited close to the coast in a context subject to the ecological influence of the open sea.

Eocene Round Herring from Monte Bolca, Italy

Fishes of the subfamily Dussumieriinae, also known as round herrings, are a small subgroup of the family Clupeidae inhabiting tropical and subtropical marine coastal waters. A new genus and species of round herring, Trollichthys bolcensis gen. et sp. nov., is described from the lower Eocene micritic limestone of Monte Bolca, northern Italy. This new clupeid taxon is based on seven partially complete articulated skeletons that exhibit a unique combination of features, including: two supramaxillae, edentulous jaw and palate bones, 41–42 preural vertebrae and 22–24 pleural ribs, pleural ribs-preural vertebrae ratio ranging 0.52–0.57, five or six supraneural bones, dorsal-fin origin located at about mid-length of the body, dorsal fin with about 16 rays, two postcleithra, pelvic-fin insertion slightly behind the dorsal-fin origin, and pelvic fin with eight rays. Trollichthys bolcensis shares several features with the extant round herring genus Spratelloides. However, because of its unique combination of features, Trollichthys bolcensis cannot be confidently assigned to any of the extant dussumieriine lineages and present evidence does not favour any particular sister-group relationship.

A reappraisal of the Eocene priacanthid fish Pristigenys substriata (Blainville, 1818) from Monte Bolca, Italy

Abstract. Priacanthids are a small family of percomorph fishes comprising fewer than 20 extant species currently assigned to four genera. One of these, Pristigenys, was established by Louis Agassiz (1835) to include the Eocene species Pristigenys substriata from Monte Bolca, and is usually regarded as a subjective senior synonym of Pseudopriacanthus. Consequently, Pristigenys currently comprises five extant species plus the fossil Pristigenys substriata. The osteology of the type species of this genus, however, is poorly known, and this makes it difficult to provide an adequate comprehensive definition of the taxonomy of the whole family. Pristigenys substriata is redescribed in detail based on five well-preserved articulated skeletons. Pristigenys substriata can be easily distinguished from other priacanthids based on its unique combination of characters. Morphological analysis of the fossil specimens reveals that there is substantial evidence to justify recognition of both Pristigenys and Pseudopriacanthus as valid genera, with extant species previously assigned to Pristigenys now referred to Pseudopriacanthus. Within the Priacanthidae, Pristigenys and Pseudopriacanthus form sister taxa and this pair can be considered as the sister-group to all remaining extant priacanthid genera (Cookeolus + [Heteropriacanthus + Priacanthus]).

Eocene sand tiger sharks (Lamniformes, Odontaspididae) from the Bolca Konservat-Lagerstätte, Italy: palaeobiology, palaeobiogeography and evolutionary significance

Abstract Here we report the first record of one of the most common and widespread Palaeogene selachians, the sand tiger shark Brachycarcharias, from the Ypresian Bolca Konservat-Lagerstätte. The combination of dental character of the 15 isolated teeth collected from the Pesciara and Monte Postale sites (e.g. anterior teeth up to 25 mm with fairly low triangular cusp decreasing regularly in width; one to two pairs of well-developed lateral cusplets; root with broadly separated lobes; upper teeth with a cusp bent distally) supports their assignment to the odontaspidid Brachycarcharias lerichei (Casier, 1946), a species widely spread across the North Hemisphere during the early Palaeogene. The unambiguous first report of this lamniform shark in the Eocene Bolca Konservat-Lagerstätte improves our knowledge concerning the diversity and palaeobiology of the cartilaginous fishes of this palaeontological site, and provides new insights about the biotic turnovers that involved the high trophic levels of the marine settings after the end-Cretaceous extinction.

A synoptic review of the Eocene (Ypresian) cartilaginous fishes (Chondrichthyes: Holocephali, Elasmobranchii) of the Bolca Konservat-Lagerstätte, Italy

Here, we review and discuss the records and taxonomy of the Ypresian (Eocene) chondrichthyans from the famous Bolca Konservat-Lagerstätte in northeastern Italy. Despite the outstanding diversity and the numerous studies focusing on the actinopterygian faunas from Pesciara and Monte Postale, the current knowledge about the systematics, taxonomy and phylogenetic relationships of the cartilaginous fishes from these Eocene sites remains elusive and largely inadequate. The celebrated Eocene Bolca Lagerstätte has yielded several exquisitely preserved articulated remains of chondrichthyan fishes in which delicate structures and soft tissues are preserved, as well as isolated teeth. The cartilaginous fish assemblage of Bolca comprises at least 17 species-level taxa belonging to 10 families in 6 orders, including selachians (Carcharhiniformes, Lamniformes), batoids (Torpediniformes, Myliobatiformes, Rajiformes) and holocephalans (Chimaeriformes). The occurrence of holocephalans represented by an isolated fin-spine of the chimeroid Ischyodus in the Bolca assemblage is reported here for the first time and represents the first record of chimeroids in the Eocene of Italy and also southern Europe. The Bolca chondrichthyan assemblage is remarkably different from those of other contemporaneous Boreal or Tethyan deposits, suggesting that its taxonomic composition is largely influenced by the palaeoenvironmental context. However, this synoptic review also highlights the importance of detailed revisions of all chondrichthyan remains from the Bolca Konservat-Lagerstätten.KurzfassungObwohl Knochenfische in der berühmten eozänen (Ypresium) Bolca-Konservatlagerstätte in NE-Italien äußerst divers sind und in den vergangenen Jahren Gegenstand zahlreicher Studien waren, ist der derzeitige Kenntnisstand über die Systematik, Taxonomie und Phylogenie der Knorpelfische aus dieser Lagerstätte überraschend gering und ungenau. In dieser Studie geben wir einen Überblick über die Knorpelfische der Bolca-Konservatlagerstätte und diskutieren ihre Verbreitung und ihre Taxonomie. Aus den beiden Fundstellen Pesciara und Monte Postale stammen einige sehr gut erhaltene Exemplare, bei denen auch sehr feine Strukturen und Weichgewebe fossilisiert sind, sowie isolierte Zähne. Die Knorpelfischfauna von Bolca umfasst mindestens 17 Arten, die zu zehn Familien und sechs Ordnungen gehören wie Haie (Carcharhiniformes, Lamniformes), Rochen (Torpediniformes, Myliobatiformes, Rajiformes) und Chimären. Chimären sind erstmals für Bolca durch einen isolierten Flossenstachel von Ischyodus belegt, was gleichzeitig der erste Nachweis für Chimären im Eozän von Italien und Süd-Europa ist. Die Knorpelfischassoziation von Bolca unterscheidet sich in ihrer taxonomischen Zusammensetzung auffällig von anderen tethyalen und auch borealen Faunen. Dies lässt vermuten, dass die taxonomische Zusammensetzung im Wesentlichen durch Umweltbedingungen bestimmt ist. Der hier präsentierte synoptische Überblick der Bolca-Knorpelfische zeigt aber auch deutlich, dass detaillierte Revisionen aller Knorpelfischreste der Bolca-Konservatlagerstätte für ein besseres Verständnis dieser wichtigen eozänen Fundstelle dringend nötig sind.

Revision of Eocene electric rays (Torpediniformes, Batomorphii) from the Bolca Konservat-Lagerstätte, Italy, reveals the first fossil embryo in situ in marine batoids and provides new insights into the origin of trophic novelties in coral reef fishes

The Eocene electric ray †Titanonarke Carvalho, 2010 from the Bolca Konservat-Lagerstätte, north-eastern Italy, is redescribed in detail based upon new material from recent excavations. This taxon exhibits a combination of features (large voids between the pectoral and the axial skeleton filled in life by electric organs, anteriorly directed fan-shaped antorbital cartilages, lack of dermal denticles, long prepelvic processes, and rounded basibranchial copula with a small caudal tab) that clearly supports its assignment to the order Torpediniformes. The analysis of new material also demonstrates that the previous apparent absence of typical narcinoid characters used to diagnose †Titanonarke was the result of taphonomic biases. †Titanonarke shares at least three synapomorphies (presence of a rostral fontanelle, low number of ribs, and rostral cartilage connected to the antorbital cartilage through lateral appendices) with the extant genera Benthobatis, Diplobatis, Discopyge and Narcine, with which it forms a clade (family Narcinidae) recognized herein as unquestionably monophyletic. Moreover, based upon a single specimen of †Titanonarke that exhibits a unique combination of morphometric and meristic features, a new species of Eocene numbfish, †T. megapterygia sp. nov., is recognized. The presence of several specimens representing different ontogenetic stages of at least two species of numbfishes suggests a close association of this taxon with shallow-water habitats corresponding to coral reefs as hypothesized for the Monte Postale palaeoenvironment. The occurrence of a fossilized marine batoid embryo is reported here for the first time. Moreover, the analysis of the gut contents suggests that the dietary adaptations of †Titanonarke can be related, at least in part, to an opportunistic strategy in the context of abundant larger foraminifera in the Monte Postale palaeobiotope, suggesting that this kind of feeding mode, known to occur in present-day reefs, already was realized 50 million years ago. http://zoobank.org/urn:lsid:zoobank.org:pub:EDD6E170-CA64-4FFB-8DD1-AED2D61D5504

Non-reef environments impact the diversification of extant jacks, remoras and allies (Carangoidei, Percomorpha)

Various factors may impact the processes of diversification of a clade. In the marine realm, it has been shown that coral reef environments have promoted diversification in various fish groups. With the exception of requiem sharks, all the groups showing a higher level of diversity in reefs than in non-reef habitats have diets based predominantly on plankton, algae or benthic invertebrates. Here we explore the pattern of diversification of carangoid fishes, a clade that includes numerous piscivorous species (e.g. trevallies, jacks and dolphinfishes), using time-calibrated phylogenies as well as ecological and morphological data from both extant and fossil species. The study of carangoid morphospace suggests that reef environments played a role in their early radiation during the Eocene. However, contrary to the hypothesis of a reef-association-promoting effect, we show that habitat shifts to non-reef environments have increased the rates of morphological diversification (i.e. size and body shape) in extant carangoids. Piscivory did not have a major impact on the tempo of diversification of this group. Through the ecological radiation of carangoid fishes, we demonstrate that non-reef environments may sustain and promote processes of diversification of different marine fish groups, at least those including a large proportion of piscivorous species.

The relationships of Gasteroclupea branisai Signeux, 1964, a freshwater double-armored herring (Clupeomorpha, Ellimmichthyiformes) from the Late Cretaceous-Paleocene of South America

Abstract The clupeomorph fish Gasteroclupea branisai Signeux, 1964 from the Late Cretaceous-Paleocene of El Molino Formation, Cayara locality, Bolivia is redescribed in detail. This South American double-armored herring is the sole member of the family Gasteroclupeidae and exhibits a unique combination of features (skull roof ornamented with fine tubercles, beryciform foramen, endopterygoid teeth, about 16–18 supraneurals, a series of 35–38 predorsal and 40 ventral scutes, median fins small and containing 12–14 rays) that clearly supports its assignment to the extinct clupeomorph order Ellimmichthyiformes. G. branisai shares a number of synapomorphies with Sorbinichthys elusivo and S. africanus (five hypurals, second hypural autogenous, distal end of the second uroneural not reaching the distal end of the first uroneural) with which it forms a monophyletic assemblage recognized herein for the first time as the new suborder Sorbinichthyoidei. The peculiar morphology of Gasteroclupea that is characterized by having an extremely deep abdominal profile, hypertrophied coracoid, and elongated pectoral fins, resembles in many aspects the body plan of extant freshwater hatchetfishes of the characiform family Gasteropelecidae, suggesting that the morphological adaptations of G. branisai can be related, at least in part, to a similar lifestyle.

REDESCRIPTION OF ‘PERLEIDUS’ (OSTEICHTHYES, ACTINOPTERYGII) FROM THE EARLY TRIASSIC OF NORTHWESTERN MADAGASCAR

The revision of the material from the Lower Triassic fossil-bearing-nodule levels from northwestern Madagascar supports the assumption that the genus Perleidus De Alessandri, 1910 is not present in the Early Triassic. In the past, the presence of this genus has been reported in the Early Triassic of Angola, Canada, China, Greenland, Madagascar and Spitsbergen. More recently, it has been pointed out that these taxa may not be ascribed to Perleidus owing to several anatomical differences. The morphometric, meristic and morphological analyses revealed a remarkable ontogenetic and individual intraspecific variation among dozens of specimens from the lower Triassic of Ankitokazo basin, northwestern Madagascar and allowed to consider the two Malagasyan species P. madagascariensis Piveteau, 1934, and P. piveteaui Lehman, 1952, as a single one and to ascribe it to the new genus Teffichthys . This new genus exhibits a unique combination of features, mainly in the skull dermal bone pattern and structure of caudal fin, that clearly support its exclusion from Perleidiformes. We also suggest to extend the use of Teffichthys for the other Early Triassic ‘ Perleidus ’ species except those from southern China.