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†Wettonius angeloi n. gen. n. sp. from the Eocene of Monte Bolca. Reconstruction of the caudal skeleton, right lateral view.
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A new sailfin veliferid fish, †Wettonius angeloi n. gen. n. sp., is described based on a single specimen from the Eocene locality of Monte Bolca in northern Italy. †Wettonius angeloi n. gen. n. sp. differs from the known Veliferidae by having a relatively short lower jaw, mandibular joint anterior to the orbit, reduced number of abdominal vertebrae...
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... Genuine Lampriformes (Verliferidae and Wettonius) and Polymixiiformes (Polymixiidae and Polyspinatus) are known from the lower Eocene (Carnevale & Bannikov, 2018;Schrøder et al., 2022), and there are also Late Cretaceous otolith records that suggest the presence of Lampriformes and Polymixiiformes of modern type (Fig. 6). These otoliths are less spectacular in morphology and well in line with extant morphotypes found in the Polymixiidae and Veliferidae. ...
The Late Cretaceous was a time of blossoming teleost diversification that came to a sudden restriction and partial termination during the extinction event at the Cretaceous–Paleogene (K/Pg) boundary. Among the dominant and diverse Late Cretaceous teleost groups prior to the K/Pg boundary event were certain pelagic Aulopiformes (e.g., Ichthyotringoidei and Enchodontoidei) and a large variety of basal Acanthomorpha whose relationships are difficult to determine. The skeletal record diminishes during the late Campanian and is low in the Maastrichtian and Paleocene, constituting the so-called ‘Patterson’s Gap’. Recent studies of fossil otoliths, however, have significantly increased the number of taxa recognised for this time inerval, but most of the putative extinct forms lack adequate calibration with otoliths found in situ in articulated skeletons. However, the otolith assemblages do confirm the presence of great morphological diversity among Aulopiformes and Acanthomorpha incertae sedis that became extinct at the K/Pg boundary. In the present review, we elucidate the effect of the K/Pg boundary from an otolith perspective and categorise extinct lineages and survivors. It is interesting to recognise that several of the surviving lineages are represented by groups that probably originated during the Late Cretaceous but were not particularly common up to the K/Pg boundary and began to expand rapidly and diversify during the early Paleogene. Such lineages probably took advantage to populate void ecospace that may have opened following the extirpation of previously dominant lineages. During the early Paleogene, the otolith record shows that the Ophidiiformes and perciforms s. lat. were the ones that diversified the most rapidly and became the most abundant, and in certain areas associated with the Gadiformes.
... The Ypresian Fur Formation of Denmark yields the enigmatic †Palaeocentrotus plus an undescribed, †Analectislike taxon (Bonde, 1966;Bonde et al., 2008). The earliest lampriforms that can be confidently classified within modern families are two veliferids ( †Veronavelifer and †Wettonius) from the late Ypresian of Bolca, Italy (Bannikov, 1990(Bannikov, , 2014Carnevale et al., 2014;Carnevale & Bannikov, 2018)--although Near et al. (2013) have argued that †Turkmene is a lamprid, without providing a detailed justification. Bolca also yields two taxa historically described as lampriforms: the enigmatic 'Pegasus' (Carnevale et al., 2014;Carnevale & Bannikov, 2018) and †Bajaichthys (Bannikov, 2014;Sorbini & Bottura, 1988), the latter now regarded as a morphologically distinctive zeiform . ...
... The earliest lampriforms that can be confidently classified within modern families are two veliferids ( †Veronavelifer and †Wettonius) from the late Ypresian of Bolca, Italy (Bannikov, 1990(Bannikov, , 2014Carnevale et al., 2014;Carnevale & Bannikov, 2018)--although Near et al. (2013) have argued that †Turkmene is a lamprid, without providing a detailed justification. Bolca also yields two taxa historically described as lampriforms: the enigmatic 'Pegasus' (Carnevale et al., 2014;Carnevale & Bannikov, 2018) and †Bajaichthys (Bannikov, 2014;Sorbini & Bottura, 1988), the latter now regarded as a morphologically distinctive zeiform . The Paleogene lampriform fossil record includes a variety of taeniosomes, all assigned to Lophotidae: †Eolophotes from the Lutetian of Georgia (Daniltshenko, 1980), †Protolophotus and †Babelichthys from the late Eocene of Iran (Arambourg, 1943;Davesne, 2017;Walters, 1957) and ...
The early Eocene fossil assemblage of the London Clay (Southeastern England) is a key window to the early Paleogene diversification of teleost fishes in the open ocean. Despite their three-dimensional preservation that offers unique insight into skeletal anatomy, the London Clay fossils are still poorly described for the most part. †Whitephippus tamensis is a fossil teleost from this assemblage, known by several well-preserved specimens. Based on a complete description of the known material, including previously hidden structures (braincase, hyoid, and branchial arches) revealed through 3D microtomography, we reinterpret †Whitephippus as an early member of the teleost group Lampriformes. More specifically, the anatomy of †Whitephippus indicates that it is likely a member of the so-called 'pelagic clade' including modern opahs and oarfishes. This redescription of †Whitephippus provides the earliest definitive evidence of lampriforms conquering the pelagic environment, alongside numerous other teleost lineages.
... We expanded the dataset of Davesne et al. (2014) as modified by Davesne et al. (2016) to include all extant taeniosome genera, the fossil taeniosomes †Babelichthys olneyi (Davesne 2017), †Eolophotes lenis (Danilit'chenko 1962), †Oligolophotes fragosus (Bannikov 1999), and †Protolophotus elami (Arambourg 1966), and the veliferid †Wettonius angeloi (Carnevale and Bannikov 2018). Character states from added operational taxonomic units (OTUs) were scored based on examination of specimens in the collections of the Muséum National d'Histoire Naturelle, Paris, France ( †Babelichthys olneyi, †Protolophotus elami), and from the literature (see Danilit'chenko 1962, Arambourg 1966, Bannikov 1999, Davesne 2017, Carnevale and Bannikov 2018. ...
... We expanded the dataset of Davesne et al. (2014) as modified by Davesne et al. (2016) to include all extant taeniosome genera, the fossil taeniosomes †Babelichthys olneyi (Davesne 2017), †Eolophotes lenis (Danilit'chenko 1962), †Oligolophotes fragosus (Bannikov 1999), and †Protolophotus elami (Arambourg 1966), and the veliferid †Wettonius angeloi (Carnevale and Bannikov 2018). Character states from added operational taxonomic units (OTUs) were scored based on examination of specimens in the collections of the Muséum National d'Histoire Naturelle, Paris, France ( †Babelichthys olneyi, †Protolophotus elami), and from the literature (see Danilit'chenko 1962, Arambourg 1966, Bannikov 1999, Davesne 2017, Carnevale and Bannikov 2018. We did not include the extinct taxa †Bathysoma lutkeni, †Palaeocentrotus boggildi, and the recently described †Oechsleria unterfeldensis (Micklich and Bannikov 2023), because all of these to our knowledge are known from one or two partial skeletons with poorly preserved cranial material, making them non-ideal for phylogenetic analysis. ...
... The topology of the strict consensus (Fig. 2) is similar to previously published phylogenies inferred using the earlier versions of the dataset (Davesne et al. 2014. Notably, our new morphological phylogenetic analysis resolves †Wettonius angeloi as a pan-veliferid and †Protolophotus elami, †Eolophotes lenis, and †Oligolophotes fragosus as crown taeniosomes, placements that were previously hypothesized on the basis of qualitative anatomical comparisons (Bannikov 1999, Davesne 2017, Carnevale and Bannikov 2018. In the morphological phylogeny (Fig. 2) leading to Regalecidae (Regalecus and Agrostichthys parkeri), Radiicephalus, and Trachipteridae (Trachipterus, Desmodema, and Zu). ...
Ray-finned fishes, which compose nearly half of living vertebrate diversity, provide an excellent system for studying the evolution of novel body forms. Lampriformes is a species-poor lineage of acanthomorph ray-finned fishes that has evolved two very different and highly specialized body plans suited to life in pelagic oceanic habitats: the deep, round-bodied bathysomes and the ribbon-like taeniosomes. Here, we present a new phylogenetic hypothesis and divergence time estimates for lampriform fishes based on an updated morphological dataset and DNA sequences from nuclear genes for all but one of the living lampriform families and 55% of recognized extant genera. Our analyses resolve two major clades in Lampriformes: the Bathysomi and the Taeniosomi. A time calibrated phylogeny shows that the origin of living lampriforms coincides with the aftermath of the Cretaceous–Palaeogene extinction and that anatomically modern pelagic morphotypes evolved 10 Myr after the start of the Palaeogene.
... The Ypresian Fur Formation of Denmark yields the enigmatic †Palaeocentrotus plus an undescribed, †Analectis-like taxon (Bonde, 1966;Bonde et al., 2008). The earliest lampriforms that can be confidently classified within modern families are two veliferids ( †Veronavelifer and †Wettonius) from the late Ypresian of Bolca, Italy (Bannikov, 1990(Bannikov, , 2014Carnevale et al., 2014;Carnevale & Bannikov, 2018)-although Near et al. (2013) have argued that †Turkmene is a lamprid, without providing a detailed justification. Bolca also yields two taxa historically described as lampriforms: the enigmatic 'Pegasus' (Carnevale et al., 2014;Carnevale & Bannikov, 2018) and †Bajaichthys (Bannikov, 2014;Sorbini & Bottura, 1988), the latter now regarded as a morphologically distinctive zeiform . ...
... The earliest lampriforms that can be confidently classified within modern families are two veliferids ( †Veronavelifer and †Wettonius) from the late Ypresian of Bolca, Italy (Bannikov, 1990(Bannikov, , 2014Carnevale et al., 2014;Carnevale & Bannikov, 2018)-although Near et al. (2013) have argued that †Turkmene is a lamprid, without providing a detailed justification. Bolca also yields two taxa historically described as lampriforms: the enigmatic 'Pegasus' (Carnevale et al., 2014;Carnevale & Bannikov, 2018) and †Bajaichthys (Bannikov, 2014;Sorbini & Bottura, 1988), the latter now regarded as a morphologically distinctive zeiform . The Paleogene lampriform fossil record includes a variety of taeniosomes, all assigned to Lophotidae: †Eolophotes from the Lutetian of Georgia (Daniltshenko, 1980), †Protolophotus and †Babelichthys from the late Eocene of Iran (Arambourg, 1943;Davesne, 2017;Walters, 1957) and †Oligolophotes from the early Oligocene of Russia (Bannikov, 1999). ...
The early Eocene fossil assemblage of the London Clay (Southeastern England) is a key window to the early Palaeogene diversification of teleost fishes in the open ocean. Despite their three-dimensional preservation that offers unique insight into skeletal anatomy, the London Clay fossils are still poorly described for the most part. Whitephippus tamensis is a fossil teleost from this assemblage, known by several well-preserved specimens. Based on a complete description of the known material, including hidden structures (braincase, hyoid and branchial arches) revealed through 3D microtomography, we reinterpret Whitephippus as an early member of the teleost group Lampriformes. More specifically, the anatomy of Whitephippus indicates that it is likely a member of the so-called "pelagic clade" including modern opahs and oarfishes. This redescription of Whitephippus provides the earliest definitive evidence of lampriforms conquering the pelagic environment, alongside numerous other teleost lineages.
... In 1999, Sorbini and Sorbini described a fossil of the oldest known lampriform fish, Nardovelifer altipinnis, found in the Cretaceous deposits of Nardò, Italy [62]. Carnevale and Bannikov [87], and Papazzone et al. [88] described in Eocene deposits from Verona, Italy, two ancient species, Bajaichthys elegans and Veronavelifer sorbinii. Carnevale [84], reported about the first fossil of the ribbonfish Trachipterus mauritanicus from a Miocene locality in northwestern Algeria. ...
... The Veliferidae family is considered to be the sister group of all other Lampriformes [61]. For this reason, despite being very rare, it remains one of the most studied families from the point of view of phylogenetic relationships [85][86][87]. ...
Lampriformes are circumglobally distributed and contain several families of strictly marine bony fishes that have a peculiar morphology. Lampriformes systematics is affected by limitations in biometric, meristic, and molecular data; for this reason, it underwent several rearrangements in the past. This review aimed to describe the biological and ecological characteristics of the order Lampriformes, summarizing the current taxonomy of the group. The main aim was to clarify what is known about the distribution of the order Lampriformes in the Mediterranean Sea, collecting all the scarce and fragmented reports and notes on their occurrence. Knowledge scarcity is due to their solitary nature, in addition to their low to absent economic value. Despite this, the order Lampriformes represents a taxon of high biological and ecological importance. The high depth range of distribution characterizes their lifestyle. In the Mediterranean Sea, four families are present—Lampridae, Lophotidae, Regalecidae, and Trachipteridae—with the following species respectively, Lampris guttatus (Brünnich, 1788), Lophotus lacepede (Giorna, 1809), Regalecus glesne (Ascanius, 1772), Trachipterus arcticus (Brünnich, 1788), T. trachypterus (Gmelin, 1789), and Zu cristatus (Bonelli, 1819). Data deficiencies affect information on this taxon; the present review, which collected all the reports of the Mediterranean Sea, creates a baseline for depicting the biogeography of these rare and important species.
... There are only three known genera and species: †Nardovelifer altipinnis Sorbini and Sorbini, 1999, from the upper Campanian-lower Maastrichtian of Apulia in Italy; this species, however, has more recently been considered to be a stem lampridiform (Delbarre et al. 2016). In addition, there are †Veronavelifer sorbinii Bannikov, 1991 and†Wettonius angeloi Carnevale andBannikov, 2018, both of which are from the Early Eocene (Late Ypresian) of Monte Bolca (also in Italy). Furthermore, otolith records from the Middle Eocene (Lutetian) of the Paris Basin were described by Nolf (1973Nolf ( , 2013 as "aff. ...
... For more detailed information concerning these taxa and other extant and fossil Lampridiformes and veliferoids, see (e.g.) Bannikov (1991), Olney et al. (1993), Bannikov (1999), Sorbini and Sorbini (1999), Bannikov (2014), Davesne et al. (2014), as well as Carnevale and Bannikov (2018). ...
... Large parts of the supraoccipital are more or less sufficiently preserved. They clearly exhibit a lateral ornamentation that consists of several antero-dorsal ridges that very much resembles the one described and figured for †Wettonius angeloi (Carnevale and Bannikov 2018). The outline of the forehead of †Oechsleria unterfeldensis seems to be rather steep and encloses an angle of about 70° with the neurocranial base. ...
These are the first and so far only records of a sailfin velifer fish from the Lower Oligocene of the Unterfeld (“Frauenweiler”) clay pit at Rauenberg (S Germany), and only the fourth fossil skeletal finds of this group worldwide. The new genus and species † Oechsleria unterfeldensis is described in detail, diagnosed, and compared to other fossil and extant representatives of the Veliferidae. It appears to be a comparatively small-sized fish, which differs from the other representatives of this family, apart from body proportions, by having a lower number of vertebrae and dorsal-fin rays, the absence of massive spines in the dorsal and anal fins, four anal-fin pterygiophores in front of the anteriormost haemal spine, and a different morphology and size of various skeletal elements. Amongst others, the following character states are of relevance: a supraoccipital that is laterally sculptured by strong surface ridges; a weak ascending process of the premaxilla that is shorter than in the other veliferid taxa; compound (fused) anteriormost dorsal- and anal-fin pterygiophores, together with spineless dorsal and anal fins with unornamented rays (no spinules), of which the majority are bilaterally paired and both halves are fused only proximally but separate and segmented distally; a short coracoid that does not reach the ventral body margin; a broad and anterodorsally directed pelvic bone that bears a well-developed postpelvic process, and probably also the presence of a short pointed neural spine (in contrast to a distally blunt one and/or a low crest) on the second preural centrum. All in all, the new fossil records remarkably increase the known biodiversity of the Veliferidae. They also expand the known palaeogeographic range of this family as far as to the Western Paratethys. With reference to the occurrence and life habits as deduced from the extant forms, they seem to be another Indo-Pacific (respectively Palaeo-Mediterranean, when referring to the fossil forms) element of the Grube Unterfeld fish fauna with a preference for deeper waters and affinities to temperate to tropical climates.
... There is only a limited fossil record of this family that consists of two Eocene species known from the Monte Bolca deposits in northern Italy. One is Veronavelifer sorbinii (Bannikov, 1991), likely closely related to Metavelifer multiradiatus, and the other is Wettonius angeloi (Carnevale and Bannikov, 2018), which may be closely related to Velifer hypselopterus. The specimen discovered at the Fisher/Sullivan site is apparently the first record of this family in the Western Hemisphere. ...
The bony fish fauna of the early Eocene Nanjemoy Formation is here expanded and revised to include 44 taxa identifiable to a family or lower taxonomic level. This bony fish fauna represents a diverse mix of taxa whose modern relatives are found in warm temperate, subtropical, and tropical climates. A few of these fish taxa are known only from eastern North America, but two-thirds of them have close affinity or identity with the bony fish fauna of the early Eocene London Clay in southern England. A lesser but clearly discernable affinity also exists with the early middle Eocene Monte Bolca fish fauna in northern Italy.
... The assemblage consists of about 250 taxa of sharks, batoids, pycnodontiforms and teleosts (e.g., Bannikov, 2014;Carnevale et al., 2014;Cawley et al., 2018;Marramà et al., 2018a) that inhabited a peri-reefal paleobiotope (see , and includes the earliest record of many reef-fish groups providing evidence of the apparent Cenozoic stability of the structure and composition of tropical and subtropical marine ichthyofaunas (Bellwood & Wainwright, 2002;Carnevale, 2006;Marramà et al., 2016b, c). In the last few decades a considerable amount of taxonomic and systematic studies has been devoted to several groups of bony and cartilaginous fishes collected at Monte Bolca, including anguilliforms (Blot, 1978(Blot, , 1984, aulopiforms , atheriniforms (Bannikov, 2008), batoids (Marramà et al., 2018b(Marramà et al., , 2019b, beryciforms (Sorbini, 1975(Sorbini, , 1984Sorbini & Tirapelle, 1975), clupeomorphs (Marramà & Carnevale, 2015a, b, 2016Marramà et al., 2019a), lampridiforms (Bannikov, 1991;Carnevale & Bannikov, 2018), lophiiforms (Carnevale & Pietsch, 2009, pharyngognaths (e.g., Bannikov & Carnevale, 2010Bannikov & Bellwood, 2017), pleuronectiforms (Chanet, 1999;Friedman, 2008), scombroids (Monsch, 2006), sparids (Day, 2003;Bannikov, 2006), syngnathiforms (Blot, 1980;Tyler, 2004;, tetraodontiforms (e.g., Tyler & Santini, 2002), zeiforms (e.g., Davesne et al., 2017), as well as of a number of other percomorphs (Blot, 1969;Bannikov & Carnevale, 2009;Carnevale et al., 2017). ...
A new callionymoid fish, †Gilmourella minuta n. gen. n. sp., is described herein based on a single specimen from the Eocene locality of Monte Bolca, northern Italy. †Gilmourella minuta n. gen. n. sp. differs from other callionymoids (families Callionymidae and Draconettidae) by having a large and elongate head, snout well-development, thin opercular bones, a preopercle with a posterior blunt spine, a ribbon-like interopercle, a subtriangular opercle with fimbriated posterior margin, an elongate and distally pointed subopercle, 18 (7+11) anteroposteriorly compact vertebrae, a caudal fin with 14 short rays, absence of the spinous dorsal-fin, dorsal and anal fins with five unbranched rays, and short pelvic fins. Within the callionymoids, †Gilmourella n. gen. shares a number of features (strongly protractile upper jaws, absence of the endopterygoid, the hypurals and parhypural consolidated into single plate, the haemal spine of penultimate vertebra fused to centrum, anal-fin rays being mostly unbranched) with the Callionymidae and together they seem to form a sister pair. In addition a brief discussion of the fossil record of the dragonets reveals that †Gilmourella minuta n. gen. n. sp. represents the oldest record for the callionymoid fishes known to date. RIASSUNTO-[Un dragoncello (Teleostei, Callionymoidei) nell'Eocene di Monte Bolca, Italia]-†Gilmourella minuta n. gen. n. sp., un nuovo callionimoide proveniente dai depositi eocenici di Monte Bolca, Italia settentrionale, viene descritto sulla base di un singolo reperto. †Gilmourella minuta n. gen. n. sp. differisce dagli altri callionimoidi (membri delle famiglie Callionymidae e Draconettidae) nel possedere la regione cefalica del corpo ampia e piuttosto allungata con un muso molto sviluppato, le ossa della serie opercolare molto sottili, il preopercolare caratterizzato posteriormente da una spina dall'apice arrotondato, l'interopercolare nastriforme, l'opercolare di forma subtriangolare con un margine posteriore fimbriato, il subopercolare allungato e appuntito distalmente, colonna vertebrale composta da 18 (7+11) vertebre compatte anteroposteriormente, pinna caudale contenente 14 brevi raggi, pinna dorsale anteriore (spinosa) assente, pinne dorsale e anale contenenti cinque raggi non biforcati distalmente e pinne pelviche brevi. Tra i callionimoidi, †Gilmourella n. gen. condivide una serie di caratteri morfologici (mascelle ampiamente protrusibili, endopterigoide assente, elementi dello scheletro caudale consolidati a formare una placca ipurale, spina emale della penultima vertebra fusa con il centro vertebrale, raggi della pinna anale in gran parte non biforcati distalmente) con i membri della famiglia Callionymidae con la quale sembra costituire un gruppo monofiletico. Una rapida analisi del registro paleontologico dei callionimoidi sembra indicare †Gilmourella minuta n. gen. n. sp. come la più antica testimonianza nota ad oggi per l'intero gruppo.
