identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03E8879FB65AFFC0FF08FA51FE60FCE0.text	03E8879FB65AFFC0FF08FA51FE60FCE0.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Clupeomorpha Greenwood, Rosen, Weitzman & Myers 1966	<div><p>Superorder Clupeomorpha Greenwood, Rosen, Weitzman &amp; Myers, 1966 (sensu Grande 1985)</p><p>TAXA INCLUDED AND RELATIONSHIPS. — † Ellimmichthyiformes + († Spratticeps + Clupeiformes).</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Skull roof ornamented (16:0&gt;1); anterior frontal fontanelle absent (24:1&gt;0); anterior arm of the orbitosphenoid short, not reaching the ethmoid region (44:1&gt;0); otophysic connection involving a diverticulum of the swim bladder that penetrates the exoccipital and extends into the prootic (48:0&gt;1); prootic bulla present (50:0&gt;1); central shaft of the maxilla long (96:1&gt;0); anterior ceratohyal short, its length less than five times its depth (144:1&gt;0); first and second postcleithra present (262:0&gt;1, 263:0&gt;1); parhypural fused with PU1 (301:0&gt;1); pelvic scute present (329:0&gt;1); prepelvic scutes present between the pelvic scute and the posterior end of the coracoid (333:0&gt;1); postpelvic scutes present (338:0&gt;1).</p><p>SUPPORT. — Relative Bremer = 88%.</p><p>REMARKS. — The superorder Clupeomorpha is strongly supported by 13 unambiguous synapomorphies and encompasses all the taxa considered in our ingroups. All the taxa can be included within two orders Clupeiformes and † Ellimmichthyiformes, containing four and ten of the families traditionally recognized, respectively. † Spratticeps is considered herein as a stem-clupeiform. Grande (1982a) divided the clade in two “Divisions”, with “Division 1” including only † Ornategulum, originally described by Forey (1973a) as a “primitive clupeomorph”, and “Division 2” including † Armigatus, his † Ellimmichthyiformes (including only † Diplomystus and † Ellimmichthys longicostatus) and the Clupeiformes . Later, Grande (1985) excluded † Ornategulum from the Clupeomorpha and included † Erichalcis in his “Division 1”. Today, the Grande’s Division 1 is invalid because † Erichalcis, originally described by Forey (1975) as a “primitive clupeomorph”, has been later proved to be a basal euteleost (Arratia 1997, 1999; Hermus et al. 2004). By excluding the Division 1 from the group, the synapomorphies of the Clupeomorpha as recognized by Grande (1985) include 1) the presence of abdominal (pelvic, prepelvic and postpelvic) scutes, 2) an otophysic connection involving a diverticulum of the swim bladder that penetrates the exoccipital and extends into the prootic within the lateral wall of the braincase, forming ossified bullae in the prootic, 3) supratemporal commissural sensory canal passing through parietals and supraoccipital, 4) second hypural fused with the first ural centrum, 5) a well-defined pre-epioccipital fossa, and 6) keeled dorsal scutes. Of these, our analysis solely recognized the first two characters as clupeomorph synapomorphies.</p><p>Although the presence of a complete series of abdominal scutes is one of the distinctive features of the clupeomorphs, the presence of taxa with a single scute (e.g., Dussumieriidae, Spratelloididae, and some engraulids), which were considered as “basal” (e.g., Whitehead 1963), resulted in a debate about the possession of a fully developed series of abdominal scutes in the most recent common ancestor (MRCA hereafter) of the clupeomorphs or whether the absence of scutes (e.g., † Ornategulum) represents a distinctive character of the basal bauplan of clupeomorphs or not, or if the taxa devoid of a complete series of abdominal scutes should be regarded as genuine clupeomorphs (see, e.g., Grande 1982a, 1985). In this perspective,our analysis supports the hypothesis that the MRCA of clupeomorphs had a complete series of abdominal scutes, which have been secondarily reduced (e.g., in Chirocentrus, round herrings, and New World anchovies) or completely lost (e.g., † Nardoclupea, † Ornategulum). Using the logical basis for morphological characters in phylogenetics (e.g., Sereno 2007) we distinguished the presence of otophysic connection, and of pterotic and prootic bullae as three distinct characters, highlighting that, beside the presence of otophysic connection involving penetration of the exoccipital by a swim bladder diverticulum, the presence of prootic bullae can be considered as a synapomorphy of the Clupeomorpha as well.</p><p>The third synapomorphy proposed by Grande (1985) (supratemporal commissural sensory canal primitively passing through parietals, or through parietals and supraoccipital) was not identified as a synapomorphy of the Clupeomorpha in our analysis for two reasons: 1) character 2 of Grande (1985) has been split in two distinct characters; 2) the supratemporal canal passing through parietals is a feature that characterizes,other than all clupeomorphs, also Chanos and † Tischlingerichthys making the state as a synapomorphy of the pair formed by Ostariophysi + Clupeomorpha, whereas the supratemporal canal passing also through the supraoccipital is a synapomorphy of the clade that includes † Spratticeps + Clupeiformes .</p><p>Also the fourth synapomorphy proposed by Grande (1985) (second hypural fused with the first ural centrum) was not identified as synapomorphy of the Clupeomorpha in our analysis, since the basalmost †ellimmichthyiform † Sorbinichthys retains the plesiomorphic teleost condition in which all the hypurals are autogenous (Bannikov &amp; Bacchia 2000; Murray &amp; Wilson 2011) and, at the same time, the condition is unknown in the stem Clupeiformes † Spratticeps (Patterson 1970) making impossible to detect the condition in the MRCA.</p><p>As far as the fifth synapomorphy proposed by Grande (1985) is concerns, although a large, well developed pre-epioccipital fossa has been detected in † Cynoclupea, † Santanaclupea, and most extant and fossil Clupeoidei, it is small or even absent in † Spratticeps, Denticeps, † Pseudoellimma, Engrauloidea and † Ellimmichthyiformes (Patterson 1970; Grande 1985; Sato 1994), thereby precluding it to represent a synapomorphy of the Clupeomorpha . The same can be said for the presence of predorsal scutes, considered as a distinctive feature of the Division 2 of Grande (1982a, 1985), since it was not identified as synapomorphy of the Clupeomorpha; although most †ellimmichthyiforms possess predorsal scutes, only a few clupeiforms possess them and their presence in key basal taxa († Spratticeps, † Santanaclupea) is unknown, making their presence a synapomorphy of the † Ellimmichthyiformes rather than of the clupeomorphs.</p><p>Interestingly,the presence of a parhypural fused with the first preural centrum (or to the compound centrum) was recovered as a key clupeomorph synapomorphy. Although Grande (1985, pp. 259-260) clearly noticed that †ellimmichthyiforms and denticipitoids have the parhypural fused to the first preural centrum, with secondary separation in clupeoids, he did not consider it as a key synapomorphy of the Clupeomorpha .</p></div>	https://treatment.plazi.org/id/03E8879FB65AFFC0FF08FA51FE60FCE0	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB65DFFC0FF1BFC3FFC64FCE1.text	03E8879FB65DFFC0FF1BFC3FFC64FCE1.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Ellimmichthyiformes Grande 1982	<div><p>Order † Ellimmichthyiformes Grande, 1982</p><p>TAXA INCLUDED AND RELATIONSHIPS. — † Sorbinichthyidae + († Codoichthys + (†Scutatuspinosidae + († Gasteroclupeidae + († Armigatidae + † Paraclupeidae))).</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Basipterygoid process of the parasphenoid present (33:0&gt;1); striations on the ventral part of opercle (201:0&gt;1); anteriormost ribs articulate with pits, followed by ribs articulating with parapophyses (220:0&gt;1); at least one predorsal scute present (318:0&gt;1).</p><p>SUPPORT. — Relative Bremer = 6%.</p><p>REMARKS. — The order † Ellimmichthyiformes was created by Grande (1982a) to contain the non-clupeiform clupeomorphs † Diplomystus dentatus and † Ellimmichthys longicostatus (but excluding † Armigatus brevissimus, † Ellimma branneri and † Ornategulum sardinioides) and defined by the lateral expansion of dorsal scute “wings” that give scutes a subrectangular shape. The same topology and diagnosis have been maintained by Grande (1985). After the discovery of new double-armored herrings, Chang &amp; Maisey (2003) include also † Armigatus, † Ellimma and † Paraclupea chetungensis (but not † Ornategulum) within the † Ellimmichthyiformes, now diagnosed by the presence of basipterygoid process of parasphenoid, beryciform foramen in anterior ceratohyal and dorsal scutes, whereas Zaragüeta Bagils (2004) diagnosed it (excluding † Armigatus, † Diplomystus and † Ornategulum) by the dorsal margin of the body with a marked angle at the insertion of the dorsal fin and prominent spine on the postpelvic scutes. Forey (2004) by including † Armigatus, † Diplomystus, † Ellimmichthys, † Paraclupea, † Sorbinichthys, and † Triplomystus proposed the presence of a S-shaped cleithrum as the only synapomorphy of the † Ellimmichthyiformes, whereas Alvarado-Ortega et al. (2008) excluded † Armigatus and diagnosed the † Ellimmichthyiformes using the presence of predorsal scutes with a subrectangular shape in the posterior part of predorsal series. More recently, Murray &amp; Wilson (2013) diagnosed the † Ellimmichthyiformes (including † Armigatus) as clupeomorphs showing medioparietal condition, frontals and parietals ornamented, basipterygoid process, beryciform foramen, predorsal scutes and three epurals; with these features, tree topology and group composition were in general retained in subsequent analyses (e.g., Murray et al. 2016; Alvarado-Ortega et al. 2020; MarramÀ &amp; Carnevale 2023). Our analysis corroborates the most recent hypotheses about the composition of the order that can be diagnosed by the presence of basipterygoid process of parasphenoid and dorsal scutes, but not by the presence of a beryciform foramen, since it is also present in basal clupeiforms (e.g., † Santanaclupea, † Scombroclupea, † Italoclupea). The presence of anteriormost ribs articulating with pits, followed by ribs articulating with parapophyses can be considered an additional unambiguous synapomorphy of the † Ellimmichthyiformes, being this pattern present in most taxa. Although the presence of endopterygoid teeth and the medioparietal condition have been often regarded as typical of the double-armoured herrings, the presence of endopterygoid teeth in several clupeiforms (e.g., Sato 1994), their absence in some †ellimmichthyiforms, and the lateroparietal condition in the basalmost †ellimmichthyiform † Sorbinichthys, suggest that these characters cannot be recognized as a synapomorphies in our hypothesis. Recent morphological and combined (morphological + molecular) analyses (Vernygora 2020; Kevrekidis et al. 2024) recovered the † Ellimmichthyiformes within the Clupeiformes, even nested within the Clupeoidei . However, as we show later, these analyses were affected by incomplete character sampling.</p></div>	https://treatment.plazi.org/id/03E8879FB65DFFC0FF1BFC3FFC64FCE1	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB65DFFC1FC21FC3CFAF7F873.text	03E8879FB65DFFC1FC21FC3CFAF7F873.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Sorbinichthyidae Bannikov & Bacchia 2000	<div><p>Family † Sorbinichthyidae Bannikov &amp; Bacchia, 2000</p><p>TAXA INCLUDED AND RELATIONSHIPS. — † Sorbinichthys elusivo + † S. africanus.</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Anterior dorsal margin of the body forming a marked angle at the dorsal-fin insertion (1:0&gt;1); lower jaw length three or more times its depth (109:0&gt;1); endopterygoid teeth present (115:0&gt;1); pleural ribs/vertebrae ratio of 0.20-0.40 (231:2&gt;0); elongated upper pectoral-fin rays (256:0&gt;1); dorsal process of the posttemporal subtriangular, wider at distal tip than at its midlength (261:0&gt;2); hypural 2 triangular, with its distal end distinctly broader than the proximal end (285:0&gt;2); caudal scutes absent (306:1&gt;0); procurrent rays deeply inserted in the interneural and interhaemal spaces (310:0&gt;1); spines on the posterior margin of the predorsal scutes (322:0&gt;1); lateral extensions of the posteriormost predorsal scutes (327:0&gt;1); 31 to 55 anal-fin pterygiophores (343:0&gt;1); anal-fin origin below the dorsal-fin base (344:0&gt;1); greatly elongated second dorsal-fin ray (353:0&gt;1).</p><p>SUPPORT. — Relative Bremer = 84%.</p><p>REMARKS. — The family † Sorbinichthyidae was created by Bannikov &amp; Bacchia (2000) to include the peculiar clupeomorph † Sorbinichthys elusivo from the Cenomanian of Namoura (Lebanon). The peculiar mix of †ellimmichthyiform (pectinate predorsal scutes, large U1, uroneural 1 autogenous) and clupeiform (beryciform foramen apparently absent, lateroparietal condition of the skull) features led the authors to leave the family as Clupeomorpha incertae sedis. Subsequent analyses (Forey 2004; Zaragüeta Bagils 2004) included † Sorbinichthys within the † Ellimmichthyiformes, with these hypotheses later corroborated by the discovery of new species ( S. africanus) that exhibits a large beryciform foramen, basipterygoid processes, and anteriormost ribs articulate with pits, followed by ribs articulating with parapophyses (Murray &amp; Wilson 2011). Other analyses placed † Diplomystus in the family † Sorbinichthyidae because they share spines on the posterior margin of predorsal scutes (Alvarado-Ortega et al. 2008). Our analysis, however, excludes the close relationship of † Diplomystus with † Sorbinichthys, because it recovered this latter sister to all the remaining †ellimmichthyiform taxa, likely due to the presence of plesiomorphic traits like lateroparietal condition, and the absence of more derived †ellimmichthyiform characters (see below), resembling some of the most recent hypotheses (e.g., Murray et al. 2016; Figueiredo &amp; Ribeiro 2017; MarramÀ et al. 2022).</p></div>	https://treatment.plazi.org/id/03E8879FB65DFFC1FC21FC3CFAF7F873	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB651FFCDFCB6FA31FE90FDE8.text	03E8879FB651FFCDFCB6FA31FE90FDE8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Armigatidae Murray & Wilson 2013	<div><p>Family † Armigatidae Murray &amp; Wilson, 2013</p><p>TAXA INCLUDED AND RELATIONSHIPS. — († Armigatus oligodentatus + († A. alticorpus + † A. dalmaticus)) + († A. elatus + ((† A. brevissimus + † A. namourensis) + († A. carrenoae + († A. plinii + † A. felixi)))).</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Incomplete series of predorsal scutes, leaving a gap posterior to occiput (319:1&gt;0); heart-shaped predorsal scutes in the anterior part of the series (320:0&gt;3); heart-shaped predorsal scutes in the posterior part of the series (321:0&gt;3).</p><p>SUPPORT. — Relative Bremer = 77%.</p><p>REMARKS. — The genus † Armigatus was established by Grande (1982a) to distinguish the fossil species † Clupea brevissima Blainville, 1818 from the Cenomanian deposits in Lebanon (previously placed in the genus † Diplomystus) from the other † Diplomystus species. The initial diagnosis of † Armigatus provided by Grande (1985) included some diagnostic features, which Forey et al. (2003) subsequently expanded. Murray &amp; Wilson (2013) later created the family † Armigatidae to include † Armigatus and † Diplomystus, though later analyses suggested this grouping was paraphyletic. Further studies have refined the diagnostic traits of † Armigatus and recognized the family † Armigatidae of Murray &amp; Wilson (2013) as paraphyletic. Although a few studies recovered † Armigatus to be a paraphyletic assemblage (e.g., Boukhalfa et al. 2018; Than-Marchese &amp; Alvarado-Ortega 2022) the most recent analyses (including our implied-weighting one) strongly support the hypothesis that both † Armigatus species can be grouped into a single monotypic family (e.g., MarramÀ &amp; Carnevale 2023; this study). The relationships between the † Armigatus species seem to agree with their age and palaeogeographical settings, supporting the hypothesis that that both the main lineages originated from an early Lower Cretaceous ancestor in the western Tethys, as the study of a new upper Hauterivian-lower Barremian species confirmed (Amalfitano et al. 2025).</p></div>	https://treatment.plazi.org/id/03E8879FB651FFCDFCB6FA31FE90FDE8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB651FFCCFED7F976FE1AF878.text	03E8879FB651FFCCFED7F976FE1AF878.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Diplomystus Cope 1877	<div><p>Genus † Diplomystus Cope, 1877</p><p>TAXA INCLUDED AND RELATIONSHIPS. — († D. dentatus + † D. shengliensis) + († D. birdi + † D. dubertreti).</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — More than 10 branchiostegal rays (153:1&gt;2); 7-9 supraneurals (208:1&gt;2); pleural rib/ preural vertebrae ratio between 0.20 and 0.40 (231:1&gt;0); spines on the posterior margin of the lateral wings of the predorsal scutes (322:0&gt;1).</p><p>SUPPORT. — Relative Bremer = 37%.</p></div>	https://treatment.plazi.org/id/03E8879FB651FFCCFED7F976FE1AF878	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB651FFCCFEF8FF1AFE52FB0B.text	03E8879FB651FFCCFEF8FF1AFE52FB0B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Gasteroclupeidae Signeaux 1964	<div><p>Family † Gasteroclupeidae (sensu MarramÀ et al. 2022)</p><p>TAXA INCLUDED AND RELATIONSHIPS. — † Guiclupea + († Gasteroclupea + † Diplomystus).</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Mouth superterminal,upward turned (2:0&gt;2); high supraoccipital crest (11:0&gt;1); posterior process of infraorbital 1 present (76:0&gt;1); quadrate-mandibular articulation anterior to orbit (131:1&gt;2); pleural ribs-preural vertebrae ratio between 0.41 and 0.46 (231:2&gt;1); proximal end of hypural 1 sharp, with no articulatory process (283:0&gt;1); 20 or more predorsal scutes (326:2&gt;0); more than 30 abdominal scutes (341:3&gt;4).</p><p>SUPPORT. — Relative Bremer = 55%.</p><p>REMARKS. — The subfamily †Gasteroclupeinae has been created by Signeaux (1964) to contain the peculiar fossil clupeomorph † Gasteroclupea from the Upper Cretaceous-Paleocene non-marine deposits of South America. Although the author recognized some affinities with the genus † Diplomystus (e.g., medioparietal condition of the skull roof, complete series of predorsal scutes with subrectangular lateral wings, etc), its general body physiognomy and the absence of pelvic fins led some authors to classify it within the Pristigasteroidea (Grande 1985; Gayet &amp; Meunier 1998). Thanks to the recent detailed redescription and, particularly, the new observations of the caudal skeleton, † Gasteroclupea has been reclassified as an †ellimmichthyiform clupeomorph with affinities with † Diplomystus and † Guiclupea, leading MarramÀ et al. (2022) to elevate the Gasteroclupeinae Signeaux, 1964 to the rank of family († Gasteroclupeidae) and containing these three genera.</p></div>	https://treatment.plazi.org/id/03E8879FB651FFCCFEF8FF1AFE52FB0B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB650FFCEFCBBFAB1FE4FFDF1.text	03E8879FB650FFCEFCBBFAB1FE4FFDF1.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Paraclupeidae Chang & Chou 1977	<div><p>Family † Paraclupeidae Chang &amp; Chou, 1977</p><p>TAXA INCLUDED AND RELATIONSHIPS. — †Thorectichthyinae + († Eoellimmichthyinae + (((“† Ellimmichthys ” spinosus + “† Ellimma ” longipectoralis) + †Triplomystinae) + † Paraclupeinae)).</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Anterior tip of the first uroneural reaching or extending anteriorly beyond the posterior margin of PU2 (298:0&gt;1); predorsal scutes increasing in size posteriorly (324:0&gt;1); posteriorly directed ventral spines on the postpelvic abdominal scutes (339:0&gt;1).</p><p>SUPPORT. — Relative Bremer = 22%.</p><p>REMARKS. — Chang &amp; Chou (1977) created the family † Paraclupeidae (= † Ellimmichthyidae of Grande 1982a) that was defined by a single character (dorsal scutes broader than long) and that accounted until then all non-clupeiform clupeomorphs († Diplomystus, † Ellimmichthys and † Paraclupea). After the recognition of † Armigatus as a separate genus (that included † Diplomystus brevissimus) by Grande (1982a), Chang &amp; Maisey (2003) emended the diagnosis of the family in order to include † Armigatus, † Diplomystus, † Ellimma, † Ellimmichthys and † Paraclupea that share the presence of laterally expanded, subrectangular dorsal scutes with some having a median keel extending posteriorly over the adjacent scute in the form of a pointed prominent spine. After the discovery of new †ellimmichthyiform genera and the inclusion of new characters, Alvarado-Ortega et al. (2008) recovered the family † Paraclupeidae as a monophyletic group that excluded † Armigatus, † Diplomystus and † Sorbinichthys but included all the other †ellimmichthyiforms that have 1) scutes of the predorsal series that increase in size posteriorly, and 2) with the posterior most ones bearing a prominent and strong spine. Subsequent studies generally agree in recovering these two synapomorphies as typically characterizing the † Paraclupeidae, although a few genera (e.g., † Codoichthys, † Scutatuspinosus) fall within or outside the family depending on the different interpretation of some states or the inclusion of new characters (e.g., Vernygora et al. 2016; Vernygora &amp; Murray 2021).</p></div>	https://treatment.plazi.org/id/03E8879FB650FFCEFCBBFAB1FE4FFDF1	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB653FFCEFC41FF15FBE2F9AA.text	03E8879FB653FFCEFC41FF15FBE2F9AA.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Eoellimmichthyinae Marramà & Carnevale 2026	<div><p>Subfamily † Eoellimmichthyinae n. subfam.</p><p>urn:lsid:zoobank.org:act:9EF46A8F-EB14-4774-AEB9-3D322A81A3EF</p><p>TAXA INCLUDED AND RELATIONSHIPS. — † Eoellimmichthys + † Ezkutuberezi .</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Preural ribs/preural vertebrae ratio of less of 0.20-0.40 (331:2&gt;0); pelvic-fin insertion anterior to the dorsal-fin origin (280:1&gt;0).</p><p>SUPPORT. — Relative Bremer = 81%.</p><p>DIAGNOSIS. — A new †paraclupeid subfamily diagnosed by the following combination of characters:body deep (60-80% SL); preanal length 67-69% SL; posterior supramaxilla with anterior process as long as the main body of the bone; preural ribs/preural vertebrae ratio of less of 0.20-0.40; dorsal process of the posttemporal subtriangular and distally sharp; last dorsal scute claw-shaped and hypertrophied, more than twice the size of the preceding scute; ascending arms of the abdominal scutes spine-like with large spaces between the wings of the adjacent scutes; pelvic-fin insertion anterior to the dorsal-fin origin; 23-25 anal-fin pterygiophores.</p><p>REMARKS. — This new subfamily is created herein to include two †paraclupeids († Eoellimmichthys, from the marine Ypresian limestone of Bolca, Italy, and † Ezkutuberezi from the upper Valanginianmiddle Barremian brackish deposit of the Spanish Basque Country) that are frequently recovered as sisters, since they share at least two unambiguous synapomorphies (see above) other than a number of similar morphological and meristic features, including a deep body (60-80% SL), posterior supramaxilla with anterior process as long as the main body of the bone, last dorsal scute claw-shaped and hypertrophied, more than twice the size of preceding scute, 23-25 anal-fin pterygiophores, and preanal length 67-69% SL (Poyato-Ariza et al. 2000; MarramÀ et al. 2019). The new subfamily can be also distinguished from the other paraclupeid subfamilies because of the presence of a subtriangular and distally sharp dorsal process of the posttemporal (vs. sub-rectangular and plate-like in other paraclupeids, except in † Thorectichthys), ascending arms of the abdominal scutes spine-like, with large spaces between the wings of adjacent scutes (vs. wide or spatulate, contacting each other for most of their length, except in † Thorectichthys). At the same time, alignment of † Eoellimmichthys and † Ezkutuberezi with the subfamily †Thorectichthyinae can be ruled out because of the absence of epipleurals in the caudal region (vs. present in † Thorectichthys) and the presence of a prominent median spine on the posteriormost predorsal scutes (vs. absent in † Thorectichthys). The † Eoellimmichthyinae n. subfam. includes taxa († Eoellimmichthys) that represent the youngest and last marine record for the † Ellimmichthyiformes (MarramÀ et al. 2019).</p></div>	https://treatment.plazi.org/id/03E8879FB653FFCEFC41FF15FBE2F9AA	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB653FFCEFEE1FD55FD7BF994.text	03E8879FB653FFCEFEE1FD55FD7BF994.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Thorectichthyinae Murray & Wilson 2013	<div><p>Subfamily † Thorectichthyinae Murray &amp; Wilson, 2013</p><p>TAXA INCLUDED AND RELATIONSHIPS. — † Thorectichthys marocensis + † T. rhadinus .</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Premaxilla long, equal or more than 50% of the maxillary length (92:0&gt;1); distal end of the second uroneural not reaching the distal end of the first uroneural (300:0&gt;1).</p><p>SUPPORT. — Relative Bremer = 47%.</p><p>REMARKS. — Murray &amp; Wilson (2013) created the genus † Thorectichthys and its own subfamily †Thorectichthyinae to include two new †ellimmichthyiform species from the late Cenomanian or early Turonian of Morocco, † T. marocensis and † T. rhadinus, the monophyly of the subfamily being supported by the absence of the median strong spine on posteriormost predorsal scutes, usually present in other paraclupeids. This is one of the reasons why this monotypic subfamily is often recovered as the basalmost †paraclupeid (e.g., Boukhalfa et al. 2018; MarramÀ et al. 2022). The genus † Thorectichthys has been recovered as paraphyletic due to a certain degree of dissimilarity of the two known species, regarding differences in the outline of the dorsal margin, skull roof ornamentation and other traits (e.g., MarramÀ &amp; Carnevale 2023). However, the present analysis corroborates the validity of the genus, as also detected by another analysis performed after the discovery of a third species, † T. fideli from the Cenomanian deposits of southeastern Mexico (Than-Marchese et al. 2024).</p></div>	https://treatment.plazi.org/id/03E8879FB653FFCEFEE1FD55FD7BF994	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB652FFCFFC28FB58FBF1F878.text	03E8879FB652FFCFFC28FB58FBF1F878.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Paraclupeinae Chang & Chou 1977	<div><p>Subfamily Paraclupeinae Chang &amp; Chou, 1977, new usage</p><p>TAXA INCLUDED AND RELATIONSHIPS. — † Ellimmichthys longicostatus + († Ellimma branneri + († Paraclupea + († Rhombichthys + († Tunisiaclupea + † Tycheroichthys)))).</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Striations on the ventral part of the opercle present (201:0&gt;1).</p><p>SUPPORT. — Relative Bremer = 57%.</p><p>REMARKS. — Chang &amp; Grande (1997) emended the diagnosis of the subfamily † Paraclupeinae to include † Paraclupea chetungensis as sister to † Ellimmichthys longicostatus based on the presence of an angle along the dorsal margin of the body at level of the dorsal-fin origin, skull roofing bones strongly sculptured with radiating ridges, supraoccipital ridge small and low, eight to nine predorsal scutes broader than long and ornamented with ridges, gap between hypural 2 and hypural 3, uroneural 2 extending to PU2. After the discovery of new †paraclupeid genera and the inclusion of new characters in phylogenetic analyses, the composition of the family has been emended to include † Triplomystus and † Tycheroichthys but excluding † Ellimmichthys (Murray &amp; Wilson 2013) . Our re-examination of the holotype of † Ellimmichthys longicostatus AMNH 734 (Fig. 3H) indicates that this taxon shows opercle striation, supporting its inclusion within this family, along with † Ellimma branneri, † Rhombichthys, † Tycheroichthys and † Tunisiaclupea that share several features and are morphologically more similar to the † Paraclupea species than to other †paraclupeids.</p></div>	https://treatment.plazi.org/id/03E8879FB652FFCFFC28FB58FBF1F878	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB652FFCFFC69FD1AFA77FBCD.text	03E8879FB652FFCFFC69FD1AFA77FBCD.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Scutatoclupea Bannikov 2015	<div><p>Genus Scutatoclupea Bannikov, 2015</p><p>TAXA INCLUDED AND RELATIONSHIPS. — † S. applegatei + † S. bacchiai.</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Pelvic fins posterior to the dorsal-fin base and closer to the caudal-fin base than to the pectoral-fin insertion (280:1&gt;2); neural spine on PU1 almost as long as the neural spine of PU2 (308:1&gt;0).</p><p>SUPPORT. — Relative Bremer = 37%.</p><p>REMARKS. — Our analysis supports the hypothesis of Bannikov (2015) that †“ Triplomystus ” applegatei Alvarado-Ortega &amp; Ovalles-Damián, 2008 can be included within the genus † Scutatoclupea .</p></div>	https://treatment.plazi.org/id/03E8879FB652FFCFFC69FD1AFA77FBCD	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB652FFCFFEF6FB02FBD6FECA.text	03E8879FB652FFCFFEF6FB02FBD6FECA.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Triplomystinae Murray & Wilson 2013	<div><p>Subfamily † Triplomystinae Murray &amp; Wilson, 2013 (new rank)</p><p>TAXA INCLUDED AND RELATIONSHIPS. — “† Triplomystus ” oligoscutatus + († Triplomystus noorae + † Scutatoclupea).</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Proximal end of the first hypural sharp, with no articulatory process (283:0&gt;1); two or one uroneurals (296:0&gt;1); postdorsal scute series present (328:0&gt;1).</p><p>SUPPORT. — Relative Bremer = 37%.</p><p>REMARKS. — Murray &amp; Wilson (2013) created the tribe †Triplomystini to include species of the genus † Triplomystus, namely † T. noorae from the Cenomanian strata of Namoura, Lebanon (Forey et al. 2003) and † T. applegatei from the Albian of Chiapas, Mexico (Alvarado-Ortega &amp;Ovalles-Damián 2008) sharing the presence of a third (postdorsal) series of scutes. After the description of † Scutatoclupea bacchiai, another Cretaceous triple-armored herring from Lebanon, Bannikov (2015) referred †“ T.” applegatei to this genus, since the Mexican species shares more features with † S. bacchiai than with † T. noorae and † T. oligoscutatus, including general body physiognomy, similar body proportions and meristic counts, relative position of the fin origins and other skeletal features. Our analysis however, recovered † Triplomystus (including † T. noorae and † T. oligoscutatus) paraphyletic due to a certain degree of dissimilarity of the two species, actually showing differences in body proportions and meristic counts, although their sister group relationship can be recovered in a non-implied weighting analysis.Due to the recurrent recovery of † Scutatoclupea and † Triplomystus as sister taxa here and in most of the previous analyses (e.g., MarramÀ &amp; Carnevale 2023) we propose to elevate the tribe †Triplomystini Murray &amp; Wilson, 2013 to the subfamily level.</p></div>	https://treatment.plazi.org/id/03E8879FB652FFCFFEF6FB02FBD6FECA	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB655FFC9FC71FA75FDA0FC2A.text	03E8879FB655FFC9FC71FA75FDA0FC2A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Clupeiformes Bleeker 1859	<div><p>Order Clupeiformes Bleeker, 1859</p><p>TAXA INCLUDED AND RELATIONSHIPS. — Denticipitoidei + († Pseudoellimma + († Leufuichthys + Clupeoidei)).</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Auditory fenestra present (23:0&gt;1); recessus lateralis present (31:0&gt;1); supratemporal system present (39:0&gt;1); vertical process of the basisphenoid absent (41:0&gt;1); pterotic bulla present (55:0&gt;1).</p><p>SUPPORT. — Relative Bremer = 59%.</p><p>REMARKS. — In the comprehensive review by Grande (1985) the order Clupeiformes comprises all the non-†ellimmichthyiform clupeomorphs grouped in two suborders, Denticipitoidei and Clupeoidei, diagnosed by the presence of the recessus lateralis, parietals separated by the supraoccipital (lateroparietal condition), and loss of the beryciform foramen (chs 9 to 11 of Grande 1985). Di Dario &amp; De Pinna (2006), recognizing the supratemporal system also in Denticeps (until then considered exclusive of the Clupeoidei) considered the presence of this character as a further synapomorphy of the Clupeiformes . Our analysis recognizes the presence of recessus lateralis and supratemporal system as synapomorphies of the Clupeiformes . The laterotemporal condition is not exclusive of the Clupeiformes, since it also occurs in Osmerus, Alepocephalus, Chanos and the basal †ellimmichthyiform † Sorbinichthys, evidencing that the MRCA of the Clupeomorpha had the lateroparietal condition (Figs 7B; 8 B-D) that has been secondarily lost in more advanced †ellimmichthyiforms. Moreover, the absence of the beryciform foramen cannot be considered a synapomorphy of the Clupeiformes as suggested by Grande (1985) since it still occurs as large perforation in some basal extinct clupeiforms († Santanaclupea), in some basal fossil clupeoids († Italoclupea and † Scombroclupea) and, although reduced in size, it is still present in some living clupeoids ( Alosa, Brevoortia, Clupea, Hyperlophus, Sardinops and Sprattus) (Maisey 1993; Sato 1994; Segura &amp; De Astarloa 2004; Taverne 2007). Interestingly, our analysis recognized the presence of the auditory fenestra, the pterotic bulla and the absence of the vertical process of basisphenoid as clupeiform synapomorphies. The auditory fenestra is a large or moderate opening in the otic region surrounded by prootic, exoccipital, and basioccipital that is likely unique of the Clupeiformes since the “auditory foramen or fenestra” present in some characiforms and osteoglossomorphs (Weitzman 1962; Hilton 2003) is likely not homologous of the auditory fenestra of the Clupeiformes (Fink &amp; Fink 1981; Mertzen 2019). Although all the clupeomorphs possess prootic bullae, the presence of pterotic bullae can be considered a synapomorphy restricted to the Clupeiformes, as they are absent in † Spratticeps and likely in all the †ellimmichthyiforms. Their absence in the extant clupeoids Clupeonella, Ethmidium and Sprattus must be therefore considered as a secondary lost (Grande 1985).</p></div>	https://treatment.plazi.org/id/03E8879FB655FFC9FC71FA75FDA0FC2A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB655FFC8FEEAF930FE1AF87A.text	03E8879FB655FFC8FEEAF930FE1AF87A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Paraclupea Sun 1956	<div><p>Genus † Paraclupea Sun, 1956</p><p>TAXA INCLUDED AND RELATIONSHIPS. — † P. chetungensis, † P. pietrarojae, † P. seilacheri, † P. solignaci .</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Supratemporal commissure sensory canal present on the supraoccipital (8:0&gt;1).</p><p>SUPPORT. — Relative Bremer = 86%.</p></div>	https://treatment.plazi.org/id/03E8879FB655FFC8FEEAF930FE1AF87A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB654FFC9FC2CFC77FB63F879.text	03E8879FB654FFC9FC2CFC77FB63F879.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Cynoclupeidae Malabarba & Di Dario 2017	<div><p>Family † Cynoclupeidae Malabarba &amp; Di Dario, 2017, new usage</p><p>TAXA INCLUDED AND RELATIONSHIPS. — † Cynoclupea + † Santanaclupea .</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Pre-epioccipital fossa large and well defined (21:0&gt;1); suspensorium posteriorly inclined (119:0&gt;1); quadrate-mandibular articulation posterior to the orbit (131:1&gt;0); 7-9 supraneurals (208:1&gt;2).</p><p>SUPPORT. — Relative Bremer = 57%.</p><p>EMENDED DIAGNOSIS. — A denticipitoid family distinguished from all the other members of the suborder by the following combination of characters: head triangular in lateral view; pre-epioccipital fossa large and well defined; snout acuminated; long and straight lower jaws with a conspicuous dentition extending well back along the oral margin of the dentary; suspensorium slightly inclined obliquely backward; quadrate-mandibular articulation posterior to the orbit; odontodes absent.</p><p>REMARKS. — The monotypic family †Cynoclupeidae was established by Malabarba &amp; Di Dario (2017) to contain the peculiar Early Cretaceous (Barremian) clupeiform † Cynoclupea nelsoni collected in the shales of the Morro do Chaves Formation of the Sergipe-Alagoas Basin in NE, Brazil, that the authors hypothesized to be closely related to the Chirocentridae and Engraulidae due to the shared presence of the dorsal tip of the symplectic in line with the dorsal margin of the quadrate, with this latter bone possessing a laminar outgrowth on its anterior margin, and conspicuous dentition in the lower jaws extending well back along the oral margin of the dentary. Our analysis, instead, suggests that † Cynoclupea is sister to † Santanaclupea silvasantoi from the late Aptian to early Albian (or late Aptian; see Melo et al. 2020) Romualdo Formation (NE Brazil) (Maisey 1993), this relationship being supported by four unambiguous synapomorphies. Since the family †Cynoclupeidae has not been formally diagnosed, we propose a new usage for the family †Cynoclupeidae, which is now diagnosed by characters shared by † Cynoclupea and † Santanaclupea, most of them actually solely used to diagnose † Cynoclupea by Malabarba &amp; Di Dario (2017).</p></div>	https://treatment.plazi.org/id/03E8879FB654FFC9FC2CFC77FB63F879	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB654FFC9FF45FBF6FBDCFCA6.text	03E8879FB654FFC9FF45FBF6FBDCFCA6.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Denticipitoidei Greenwood, Rosen, Weitzman & Myers 1966	<div><p>Suborder Denticipitoidei Greenwood, Rosen, Weitzman &amp; Myers, 1966, new usage</p><p>TAXA INCLUDED AND RELATIONSHIPS. — †Cynoclupeidae + Denticipitidae .</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Maxillo-premaxillary gap absent (88:0&gt;1); lower jaw length three or more times its depth (109:1&gt;2); pleural ribs/preural vertebrae ratio of 0.41-0.46 (231:2&gt;1); two or one uroneurals (296:0&gt;1); ascending arms of the pelvic scute short (331:1&gt;0); ascending arms of the prepelvic scutes between the pelvic scute and the posterior edge of the coracoid short (337:1&gt;0); ascending arms of the prepelvic and postpelvic scutes semicircular to subrectangular, contacting with those of adjacent scutes for almost their entire length (340:0&gt;2).</p><p>SUPPORT. — Relative Bremer = 23%.</p><p>EMENDED DIAGNOSIS. — Clupeiform fishes (having recessus lateralis and medioparietal condition) retaining the following plesiomorphic features: first ural centrum large, its size approximately similar (or slightly less) as that of the first preural centrum (vs. U1 reduced in clupeoids); first hypural articulated to the first ural centrum through massive proximal head forming a well-developed articulatory process (vs. hypural 1 not reaching the first ural centrum and having sharp proximal head, with no articulatory process); parhypural fused to PU1 (vs. autogenous); first uroneural autogenous (vs. fused to PU1, forming the pleurostyle); two or one uroneural (vs. three); complete lateral line present (vs. mostly or totally absent); keeled abdominal scutes with short or reduced semicircular to subrectangular ascending arms.</p><p>REMARKS. — The suborder Denticipitoidei was introduced by Greenwood et al. (1966) to contain the family Denticipitidae Clausen, 1959 represented by extant freshwater denticle herring Denticeps clupeoides and the Middle Eocene † Palaeodenticeps tanganikae (Greenwood 1960, 1968). Although the family and both the genera have been formally diagnosed, the suborder Denticipitoidei appears to lack formal diagnosis in its original description by Greenwood et al. (1966), although it was based on the presence of plesiomorphic morphological traits that distinguish Denticeps and † Palaeodenticeps from the other members of the Clupeiformes (i.e., the Clupeoidei), whose morphological distinctiveness has been instead satisfactory provided (see Grande 1985). In this perspective, although the suborder Denticipitoidei has not been explicitly diagnosed with unique characters, Greenwood et al. (1966: 359) provided a list of morphological characters that support its difference with respect to the other clupeiforms, and that can be used here to provide a formal diagnosis of the suborder, including features of the caudal skeleton and lateral line system, whereas the presence of odontodes (denticles) and reduction of the number of uroneurals to a single one, used traditionally since Grande (1985) to define the Denticipitoidei, are instead used herein as distinctive features of the family Denticipitidae, rather than the whole suborder. Since the features listed by Greenwood et al. (1966) are shared with the basal clupeiforms † Cynoclupea and † Santanaclupea, that our analysis recognized as closely related to the Denticipitidae to form a clade, we propose the inclusion of these two taxa within the suborder providing herein a formal diagnosis.</p></div>	https://treatment.plazi.org/id/03E8879FB654FFC9FF45FBF6FBDCFCA6	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB657FFCBFC71FAF6FD12FBCD.text	03E8879FB657FFCBFC71FAF6FD12FBCD.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Clupeoidei Bleeker 1859	<div><p>Suborder Clupeoidei Bleeker, 1859 (sensu Greenwood et al. 1966)</p><p>TAXA INCLUDED AND RELATIONSHIPS. — † Nolfia + ((† Italoclupea + † Scombroclupea) + ((Pan-Engrauloidea + ( Pristigasteroidea + Clupeoidea)))).</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Third postcleithrum present (264:0&gt;1); second postcleithrum rod-like (265:0&gt;1); fusion of first uroneural and first ural centrum, forming a pleurostyle (297:0&gt;1); three epurals (304:1&gt;0).</p><p>SUPPORT. — Relative Bremer = 69%.</p><p>REMARKS. — The Clupeoidei is the most diverse clupeiform suborder, sister to the Denticipitoidei, with its monophyly supported by several morphological characters (Grande 1985), including: 1) fusion of the first uroneural with the first preural centrum, forming a pleurostyle; 2) reduction in size of the first ural centrum; 3) loss of lateral line scales; and 4) autogenous parhypural. Patterson (1970) suggested the presence of the supratemporal system as a synapomorphy of the Clupeoidei but Di Dario &amp; De Pinna (2006), recognizing the supratemporal system also in Denticeps, and considered this character a synapomorphy of the Clupeiformes . The inclusion of some basalmost clupeiforms in our analysis highlights for the first time the gradual achievement of the traits that characterize the extant clupeoids. † Pseudoellimma lacks three of the clupeoid characters and the absence of lateral line scales cannot be checked due to incompleteness (Figueiredo 2009a), so it cannot be considered as a clupeoid. Although the separation of parhypural from PU1 was already achieved by † Leufuichthys, this taxon lacks at least one of the clupeoid synapomorphies (the pleurostyle), whereas the size of the first ural and absence of lateral line scales cannot be detected. Because of the impossibility to detect the state of these latter characters in a key taxon like † Leufuichthys, our analysis was not able to recognize them as synapomorphies of the Clupeoidei . Separation of parhypural from PU1 cannot be detected as synapomorphy of the Clupeoidei because it was already achieved in a non-clupeoid clupeiform († Leufuichthys). † Nolfia riachuelensis Figueiredo, 2009, a clupeiform species from the Albian Taquari Member of the Riachuelo Formation, in the Sergipe-Alagoas Basin (NE Brazil) is the basalmost clupeiform in which three out of four clupeoid synapomorphies can be observed (Figueiredo 2009b). Although the loss of lateral line scales cannot be checked in this species due to inadequate preservation, this synapomorphy can be inferred, as it was detected in the congeneric species † Nolfia kwangoensis Taverne, 1976 from the Cenomanian deposits of the Democratic Republic of Congo (Taverne 1997). In this perspective, it is possible to consider † Nolfia as the basalmost clupeoid. Figueiredo (2009b) considered † Nolfia as member of the Clupeidae mainly based on the presence of two rod-like postcleithra, the only character used by Grande (1985) to diagnose this family. However, recent studies recovered neither molecular nor morphological support for the monophyly of the Clupeidae sensu Grande (1985) . Nelson (1967b) suggested that the absence of abdominal scutes might be the ancestral condition within the Clupeoidei (as he considered the Dussumieriidae to be a “primitive” group within the family Clupeidae). Our analysis discards this hypothesis and is more consistent with Lavoué et al. (2014), suggesting that the MRCA of the Clupeoidei had a complete series of abdominal scutes and that the reduction in their number has been secondarily achieved at least twice within this suborder.</p></div>	https://treatment.plazi.org/id/03E8879FB657FFCBFC71FAF6FD12FBCD	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB657FFCAFF3CFF15FD7CFAB5.text	03E8879FB657FFCAFF3CFF15FD7CFAB5.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Denticipitidae Clausen 1959	<div><p>Family Denticipitidae Clausen, 1959</p><p>TAXA INCLUDED AND RELATIONSHIPS. — Denticeps + † Palaeodenticeps.</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Odontodes present (3:0&gt;1); ornamentation of the skull roof absent (16:1&gt;0); exoccipital expanded, reaching posteriorly to the level of the basioccipital (47:0&gt;1); anterior and posterior supramaxillae absent (83:1&gt;0; 84:0&gt;1); posterior arm of the preopercle present (204:0&gt;1); gap between the posteriormost ossified supraneural and the first dorsal-fin pterygiophore (217:0&gt;1); halves of neural arches of most abdominal vertebrae fused into a spine (219:0&gt;1); second postcleithrum absent (263:0&gt;1); pelvic-fins anterior to the dorsal-fin origin (280:1&gt;0); hypural 2 triangular, its distal end distinctly broader than the proximal end (285:1&gt;0); hypural diastema small, in form of a concave notch (289:0&gt;1); ascending arms of the prepelvic scutes along the ventral edge of the coracoid absent or inconspicuous (335:0&gt;1); anal-fin origin anterior to the dorsal-fin origin (344:0&gt;2); dorsal fin displaced posteriorly on the body (348:1&gt;2).</p><p>SUPPORT. — Relative Bremer = 73%.</p><p>REMARKS. — Recent studies have questioned the monophyly of Clupeiformes, as Denticeps clupeoides, the sole extant member of the Denticipitoidei,has been variously placed within the Ostariophysi or as a sister group to the clade formed by Ostariophysi + Clupeoidei (e.g., Li &amp; Ortí 2007; Bloom &amp; Lovejoy 2014; Bloom &amp; Egan 2018), although bootstrap or posterior probability values were low and arrangements likely resulting from nucleotide compositional bias (Li &amp; Ortí 2007). Conversely, evidence supporting the inclusion of Denticeps within the Clupeiformes are more robust and comes from mitogenomic (Lavoué et al. 2007, 2013; Near et al. 2012; Wang et al. 2022) and morphological data (Grande 1985; Di Dario 2004b; Di Dario &amp; De Pinna 2006), these latter including the presence of the recessus lateralis,pterotic bullae, supratemporal system and hypural 2 fused with the second ural centrum. Although we recovered the †Cynoclupeidae sisters of the Denticipitidae, our analysis still supports the traditional arrangement of the Clupeiformes since the clade formed by these families ( Denticipitoidei) is still recovered sister to the pan-Clupeoidei († Pseudoellimma + († Leufuichthys + Clupeoidei)), and therefore, within the Clupeiformes .</p></div>	https://treatment.plazi.org/id/03E8879FB657FFCAFF3CFF15FD7CFAB5	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB649FFD0FC57FBD6FE7BFBCB.text	03E8879FB649FFD0FC57FBD6FE7BFBCB.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Engrauloidea Nelson 1970	<div><p>Superfamily Engrauloidea Nelson, 1970, new usage</p><p>TAXA INCLUDED AND RELATIONSHIPS. — † Nardoclupea + ( Dussumieriidae + ( Chirocentridae + Engrauloidae n. epifam.)).</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Anterior lateral process of mesethmoid for articulation of palatine absent (63:1&gt;0); premaxillae subrectangular (90:0&gt;1); central shaft of the maxilla short, almost equal or shorter than the length of the articular head (96:0&gt;1); posterior blade of the maxilla approximately of the same size or slender than the central shaft (97:0&gt;1); dorsal and ventral profile of the maxilla nearly straight (98:0&gt;1); ectopterygoid teeth present (132:0&gt;1); more than 10 branchiostegal rays (153:1&gt;2); parapophyses on the first vertebra absent (238:1&gt;0); basal segments of the upper middle caudal-fin ray spatulate (317:0&gt;1).</p><p>SUPPORT. — Relative Bremer = 56%.</p><p>REMARKS. — This clade includes the peculiar extinct “round herring” † Nardoclupea grandei Taverne, 2002 from the Upper Cretaceous (Campanian) of Nardò (Lecce Province, southern Italy) as sister to the remaining engrauloids. Taverne (2002) described † Nardoclupea as a large-sized dussumieriine having more affinities with Dussumieria and Etrumeus than with Jenkinsia and Spratelloides . The re-examination and inclusion of this taxon in a phylogenetic context makes † Nardoclupea and the Dussumieriidae ( Etrumeus + Dussumieria) successive sister of the clade formed by the Chirocentridae + the total group Engraulidae, highlighting a step-by-step achievement of traits in the evolution of the Engrauloidea new usage.</p><p>After the introduction of the superfamily Engrauloidae n. epifam. by Nelson (1970), Grande (1985) renamed the clade as Engrauloidea to standardize the suffix (-oidae is used at epifamilial level) rather than reflect a fundamental change in the taxonomic composition of the group. The superfamily was originally created to contain the family Engraulidae Gill, 1861 with two subfamilies ( Coiliinae and Engraulinae) characterized by five traits of the branchial arches: 1) reduction of the basihyal and loss of its ossification center; 2) reduction and loss of the basihyal tooth plate; 3) elongation of the second basibranchial; 4) elongation of the dorsal parts of the gill arches; 5) fusion of tooth plates with the first and third basibranchials. Grande (1985) added two new skeletal traits: 6) suspensorium inclined obliquely backward; and 7) mesethmoid projecting anterior to vomer making the snout pig-like. Later, Di Dario (2009) suggested that the monotypic Chirocentridae ( Chirocentrus) is more closely related to the Engraulidae than to the Clupeidae, and therefore proposed a new classification of the Engrauloidea, with the inclusion of the Chirocentridae based on the presence of seven characters (see remarks of node 57), this latter hypothesis being supported by molecular and morphology-based analyses (Di Dario 2004b; Bloom &amp; Lovejoy 2014; Bloom &amp; Egan 2018; Bloom et al. 2018; Vernygora 2020; Kevrekidis et al. 2024). The close affinity of representatives of the Dussumieriini of Grande (1985) with Chirocentrus is not unusual, as molecular analyses often recovered the close relationship of Chirocentrus with Dussumieria (Wang et al. 2022), Etrumeus (Lavoué et al. 2013), or with the pair Jenkinsia + Spratelloides (Li &amp; Ortí 2007; Milec et al. 2022), or, alternatively, nested within a clade formed by Etrumeus + ( Chirocentrus + ( Jenkinsia + Spratelloides)) (Lavoué et al. 2017 a, 2019). In this perspective, we propose the inclusion of Etrumeus and Dussumieria into the superfamily Engrauloidea of Di Dario (2009) also because this relationship is still recovered if fossil taxa are excluded from the analysis (Fig. 41). Moreover, since Dussumieria and Etrumeus are found to be sister taxa also when fossil taxa are excluded and are sufficiently unique in respect to the other engrauloid families (i.e., Chirocentridae, Engraulidae, Spratelloididae) they can be included in a taxonomic rank equivalent to that of the other established families within the Engrauloidea, the Dussumieriidae Gill, 1861 .</p></div>	https://treatment.plazi.org/id/03E8879FB649FFD0FC57FBD6FE7BFBCB	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB649FFD4FEC4F8F0FBCBFC5F.text	03E8879FB649FFD4FEC4F8F0FBCBFC5F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Spratelloididae Jordan 1925	<div><p>Family Spratelloididae Jordan, 1925 (sensu Bloom &amp; Egan 2018)</p><p>TAXA INCLUDED AND RELATIONSHIPS. — Jenkinsia + Spratelloides .</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Anterior arm of the orbitosphenoid long, reaching the lateral ethmoid (44:0&gt;1); posterior process of lateral ethmoid present (65:0&gt;1); quadrate-metapterygoid fenestra present (126:0&gt;1); dorsal process of posttemporal subtriangular, wider at distal tip than at its midlength (261:0&gt;2); second and third postcleithra absent (263:1&gt;0; 264:1&gt;0); hypural 4 approximately symmetrical to the hypural 1 (292:0&gt;1); neural spine of the first preural centrum almost as long as the neural spine of PU2 (308:1&gt;0); fusion between most of the posterior margin of PU1 neural spine and the anterior margin of first uroneural (309:0&gt;1).</p><p>SUPPORT. — Relative Bremer = 73%.</p><p>REMARKS. — The small round herrings Jenkinsia and Spratelloides have been traditionally recognized to form either a subfamily of the family Dussumieriidae (the Spratelloidinae of Whitehead 1963) or a tribe within the subfamily Dussumieriinae (the Spratelloidini of Grande 1985). In both cases, their alignment with the round herring Dussumieria and Etrumeus was due to the presence of a single unkeeled W-shaped pelvic scute, along with the absence of pre- and post-pelvic scutes (Whitehead 1963; Grande 1985). We therefore follow the taxonomic ranking proposed by Bloom &amp; Egan (2018) and consider the Spratelloidinae/-ini as a family-level lineage. The close relationship between Spratelloididae or pan-Spratelloididae and the rest of the Engrauloidea is not recovered if fossil taxa are excluded from the analysis, but the Spratelloididae is instead recovered as sister to the living Clupeoidei, resembling the tree topologies of recent molecular analyses (see Remarks of node 52). Once again, this might suggest that the topologies recovered in molecular analysis (based on living taxa alone) might be biased by the lack of fossil data.</p></div>	https://treatment.plazi.org/id/03E8879FB649FFD4FEC4F8F0FBCBFC5F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB64DFFD0FC74FF15FB72FDB8.text	03E8879FB64DFFD0FC74FF15FB72FDB8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dussumieriidae Gill 1861	<div><p>Family Dussumieriidae Gill, 1861</p><p>TAXA INCLUDED AND RELATIONSHIPS. — Etrumeus + Dussumieria .</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Anterior arm of the orbitosphenoid long, reaching the lateral ethmoid (44:0&gt;1); dorsal medial lamella on orbitosphenoid present (45:0&gt;1); posterior process of lateral ethmoid present (65:0&gt;1); three or more branchiostegal rays on posterior ceratohyal (156:2&gt;3); ossified occipital epineurals present (233:0&gt;1).</p><p>SUPPORT. — Relative Bremer = 51%.</p></div>	https://treatment.plazi.org/id/03E8879FB64DFFD0FC74FF15FB72FDB8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB64CFFD1FC10FF15FBC1FC56.text	03E8879FB64CFFD1FC10FF15FBC1FC56.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Clupeopsidae Marramà & Carnevale 2026	<div><p>Family Clupeopsidae n. fam.</p><p>urn:lsid:zoobank.org:act:EF701463-967A-4F09-81DA-1AAC0D614B47</p><p>TAXA INCLUDED AND RELATIONSHIPS. — † Clupeopsis + † Monosmilus .</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Anterior arm of orbitosphenoid long, reaching the lateral ethmoid (44:0&gt;1); posterior process of lateral ethmoid present (65:0&gt;1); vomerine teeth massive and fang-like (70:0&gt;1); maxillary teeth absent (103:1&gt;0); endopterygoid teeth absent (115:1&gt;0); ectopterygoid teeth absent (132:1&gt;0); lateral horizontal lamina of the ectopterygoid present (136:0&gt;1).</p><p>SUPPORT. — Relative Bremer = 81%.</p><p>DIAGNOSIS. — A new engrauloid family whose representatives are characterized by a unique autapomorphy (lateral horizontal lamina of the ectopterygoid that overlies the maxilla) and the following combination of characters: a single massive, enlarged vomerine tooth (fang); single row of enlarged dentary fangs; maxilla edentulous; endopterygoid and ectopterygoid toothless.</p><p>REMARKS. — Since our analysis corroborates the hypothesis of Capobianco et al. (2020) about the sister group relationships between † Clupeopsis and † Monosmilus, we propose the creation of a new family diagnosed by the combination of characters recovered by our parsimony analysis and the features already evidenced by Capobianco et al. (2020).</p></div>	https://treatment.plazi.org/id/03E8879FB64CFFD1FC10FF15FBC1FC56	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB64CFFD1FC08FBD6FAA4F87E.text	03E8879FB64CFFD1FC08FBD6FAA4F87E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Engraulidae Gill 1861	<div><p>Family Engraulidae Gill, 1861</p><p>TAXA INCLUDED AND RELATIONSHIPS. — Coiliinae + pan-Engraulinae.</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Pre-epioccipital fossa reduced or obliterated (21:1&gt;0); four discrete openings present in the otic region:the opening for the infraorbital sensory canal opens between sphenotic, pterotic and frontal;the preopercular sensory canal and the cephalic sensory canal open on the pterotic; the accessory temporal canal opens on the pterotic and frontal (32:0&gt;1); anterior ceratohyal length five times or more than its dept (144:0&gt;1).</p><p>SUPPORT. — Relative Bremer = 69%.</p><p>REMARKS. — Anchovies constitute a monophyletic group (the Engraulidae) that is traditionally divided in two distinct subgroups: the subfamily Coiliinae (=Coiliidae or Old World anchovies of Grande &amp; Nelson 1985), including Coilia, Lycothrissa, Papuengraulis, Setipinna and Thryssa, and the subfamily Engraulinae (= Engraulidae of Grande &amp; Nelson 1985) including Stolephorus, Encrasicholina, Engraulis; and the New World anchovies ( Anchoa, Anchovia, Anchoviella, Cetengraulis, Lycengraulis, Pterengraulis). The family and both the subfamilies are recurrently recovered as monophyletic in morphological and molecular analyses, and our analysis corroborates this assumption. Nelson (1970) provided five diagnostic characters to distinguish anchovies from other clupeoid fishes: 1) reduction of the basihyal and loss of its ossification center; 2) reduction and loss of the basihyal tooth plate; 3) elongation of the second basibranchial; 4) elongation of the dorsal parts of the gill arches; 5) fusion of tooth plates with the first and third basibranchials. Grande (1985) added two further skeletal traits: 6) suspensorium inclined obliquely backward; and 7) mesethmoid projecting in advance of vomer making the snout pig-like. Stephens (2010) evidenced the role of an expanded orbital reticulum of the cephalic lateralis system as potential further synapomorphy of the Engraulidae .However, since it cannot be detected in † Clupeopsis or † Monosmilus, our analysis was not able to detect it as synapomorphy of the group.</p></div>	https://treatment.plazi.org/id/03E8879FB64CFFD1FC08FBD6FAA4F87E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB64CFFD1FECEFDA0FE6EF878.text	03E8879FB64CFFD1FECEFDA0FE6EF878.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Engrauloidae Marramà & Carnevale 2026	<div><p>Epifamily Engrauloidae n. epifam.</p><p>TAXA INCLUDED AND RELATIONSHIPS. — Clupeopsidae n. fam. + Engraulidae .</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Lateral expansion of the frontal bone above the orbit present (13:0&gt;1); pterotic bulla greatly expanded dorsally, obliterating the pre-epioccipital fossa (56:0&gt;1); dorsal process of the mesethmoid projecting in advance of the vomer (61:0&gt;2); vomerine teeth grouped into two well-defined groups,each at anterolateral end of the vomer (71&gt;0); suspensorium inclined obliquely backward (119:0&gt;1); articulation between the ventral limb of the hyomandibula and quadrate present (124:0&gt;1); dorsal tip of the symplectic ventral to the proximal margin of the quadrate (130:1&gt;2); quadrate-mandibular articulation posterior to the orbit (131:1&gt;0); basihyal reduced or absent (170:1&gt;0).</p><p>SUPPORT. — Relative Bremer = 82%.</p><p>DIAGNOSIS. — A new engrauloid epifamily diagnosed by the following unique combination of characters: lateral expansion of the frontal bone above the orbit; pterotic bulla greatly expanded dorsally, obliterating the pre-epioccipital fossa; dorsal process of the mesethmoid projecting in advance of the vomer; vomerine teeth grouped into two well-defined groups, each at anterolateral end of the vomer; suspensorium inclined obliquely backward; articulation between the ventral limb of the hyomandibula and quadrate; dorsal tip of the symplectic ventral to the proximal margin of the quadrate; quadrate-mandibular articulation posterior to the orbit; basihyal reduced or absent.</p><p>REMARKS. — Capobianco et al. (2020) redescribed in detail the Ypresian stem-engraulid † Clupeopsis straeleni Casier, 1946 and described the Lutetian † Monosmilus chureloides Capobianco et al., 2020 from Pakistan, bases on three-dimensional fossils. A number of derived features (e.g., suspensorium posteriorly inclined, mesethmoid projecting in advance of the vomers, substantial portion of the metapterygoid situated anterodorsal to the quadrate, ventral limb of the hyomandibula meeting the posterior margin of the quadrate and absence of the bony basihyal and basihyal tooth plate) support their close affinity with the Engraulidae, leading their inclusion into the Engrauloidea . However, † Clupeopsis and † Monosmilus lack at least one of the engraulid synapomorphies listed by Nelson (1970) (i.e., first basibranchial and basibranchial tooth plate not fused) plus other typical engraulid features (anteriorly - rather than posteriorly - directed articular surface of the quadrate) that exclude them from the crown Engraulidae (Capobianco et al. 2020) . Since our analysis corroborates the hypotheses proposed by Capobianco et al. (2020), we propose to include these two taxa and the Engraulidae in their own epifamily, the Engrauloidae n. epifam., that can be diagnosed by the characters recovered by our analysis.</p></div>	https://treatment.plazi.org/id/03E8879FB64CFFD1FECEFDA0FE6EF878	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB64FFFD2FECFFF15FE7CFB4E.text	03E8879FB64FFFD2FECFFF15FE7CFB4E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Coiliinae Bleeker 1872	<div><p>Subfamily Coiliinae Bleeker, 1872 (sensu Grande &amp; Nelson 1985)</p><p>TAXA INCLUDED AND RELATIONSHIPS. — Lycothrissa + ( Thryssa + ( Setipinna + Coilia)).</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Basihyal teeth or tooth plate absent (171:1&gt;0); first mediopharyngobranchial cartilage present (192:0&gt;1); Baudelot’s ligament originates at both first and second vertebra (235:1&gt;2); Baudelot’s ligament not ossified (236:1&gt;0); hypural diastema absent (288:0&gt;1); hypurals 5 and 6 not in line with the lower hypurals (projecting more posteriorly) (293:0&gt;1); peg on the lower middle caudal-fin ray absent (314:1&gt;0); at least one predorsal scute present (318:0&gt;1); postpelvic scutes present (338:0&gt;1); 10-20 abdominal scutes (341:1&gt;2).</p><p>SUPPORT. — Relative Bremer = 69%.</p><p>REMARKS. — Whitehead (1972) divided the Engraulidae into two subfamilies, the Engraulinae and the monotypic Coiliinae, this latter including Coilia only, to separate the peculiar rat-tailed anchovy from the other members of the family. Grande (1985) considered this division as artificial because most of the Old World anchovy genera ( Lycothrissa, Thryssa, Papuengraulis, Setipinna) are more closely related to Coilia than to other anchovies, as also suggested by Jordan &amp; Seale (1926). Grande (1985) and Grande &amp; Nelson (1985) tentatively assumed that the Coiliinae form a distinct clade from the other anchovies, with their monophyly supported by at least three morphological features: 1) loss of the peg on the lower middle caudal-fin ray, 2) posterior margin of the fifth and sixth hypurals not in line with that of the lower elements of the series, and 3) absence (or remarkable reduction) of the posteroventral notch in the third hypural. The monophyly of the Coiliinae tentatively assumed by Grande &amp; Nelson (1985) was later confirmed by several molecular studies (e.g., Lavoué et al. 2010, 2013; Wang et al. 2018, 2022), other than by our analysis that was able to recover six additional unambiguous synapomorphies supporting the clade.</p></div>	https://treatment.plazi.org/id/03E8879FB64FFFD2FECFFF15FE7CFB4E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB64FFFD3FC7AFB36FE81FE51.text	03E8879FB64FFFD3FC7AFB36FE81FE51.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Engraulinae Gill 1861	<div><p>Subfamily Engraulinae Gill, 1861 (sensu Grande &amp; Nelson 1985)</p><p>TAXA INCLUDED AND RELATIONSHIPS. — Stolephorus + ( Encrasicholina + Engraulini)</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Gap in the tooth row of the dentary present (107:0&gt;1); anteriormost two supraneurals in close proximity (215:0&gt;1); anteriormost two supraneurals having different shape respect to the others (216:0&gt;1); keel of the pelvic scute poorly developed or inconspicuous (332:1&gt;0).</p><p>SUPPORT. — Relative Bremer = 64%.</p><p>REMARKS. — Nelson (1983) provided osteological evidence (ramifications of preopercular and temporal sensory canals; first epibranchial with fused tooth plate; caudal skeleton with first preural and first ural centra) that the New World anchovies plus the cosmopolitan Engraulis and the Indo-Pacific Encrasicholina form a well-defined monophyletic group. Grande (1985) expanded the list of synapomorphies of this group, adding the loss of the dorsal scutes (although still present in some species of Stolephorus). Grande &amp; Nelson (1985) further demonstrated that the genus Stolephorus is the closest relative to this assemblage, jointly forming the subfamily Engraulinae ( Engraulidae of Grande &amp; Nelson 1985), because all of them can be diagnosed by the presence of oval eggs, nearly complete spine on the first preural centrum, two epurals, rear edge of hypurals 5 and 6 in line with the lower hypurals, close positioning of the first two supraneurals, fusion of ecto- and metapterygoid, gap in the tooth row of the dentary. Stolephorus and Encrasicholina were supposed to be successive sisters to the New World genera plus Engraulis, since they still retain less than nine needle-like prepelvic scutes with reduced lateral wings, which are absent in the remaining engrauline genera. Our analysis corroborates this hypothesis, similarly to several molecular analyses (Bloom &amp; Lovejoy 2012; Lavoué et al. 2017b; Bloom &amp; Egan 2018).</p></div>	https://treatment.plazi.org/id/03E8879FB64FFFD3FC7AFB36FE81FE51	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB64EFFD3FEAFFB36FE63F87B.text	03E8879FB64EFFD3FEAFFB36FE63F87B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Engraulini Gill 1861	<div><p>Tribe Engraulini (sensu Grande &amp; Nelson 1985)</p><p>TAXA INCLUDED AND RELATIONSHIPS. — Lycengraulis + ( Anchoviella + (( Anchovia + Anchoa) + ( Pterengraulis + ( Cetengraulis + Engraulis)))).</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Posterior frontal fontanelles closed in adult individuals (28:1&gt;0); contralateral paired infrapharyngobranchials close to each other (185:0&gt;1); prepelvic scutes between the pelvic scute and the posterior end of the coracoid absent (333:1&gt;0); single abdominal scute (341:1&gt;0).</p><p>SUPPORT. — Relative Bremer = 51%.</p><p>REMARKS. — This tribe includes Engraulis and the so called New World anchovies ( Anchoa, Anchovia, Anchoviella, Cetengraulis, Lycengraulis, Pterengraulis) that share the absence of abdominal scutes, except for the pelvic scute just in front of pelvic fins (Jordan &amp; Seale 1926; Grande &amp; Nelson 1985). Although this group is recurrently recovered as monophyletic in several morphological and molecular analyses, there is usually no con sensu s about the intrarelationships of the taxa included within the clade [compare, e.g., Di Dario (2004b), Vernygora (2020) and this work with, e.g., Bloom &amp; Lovejoy (2012), Bloom &amp; Egan (2018), Bloom et al. (2018)]. Due to limited sampling of Engraulini species, our analysis cannot test the hypothesis of Bloom &amp; Lovejoy (2012) about the separation of the so called “marine clade” from the “freshwater clade” since we included only marine species, except for a single freshwater taxon ( Pterengraulis atherinoides).</p></div>	https://treatment.plazi.org/id/03E8879FB64EFFD3FEAFFB36FE63F87B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB641FFDCFF3DF9B1FBF5F878.text	03E8879FB641FFDCFF3DF9B1FBF5F878.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Pristigasteridae Bleeker 1872	<div><p>Family Pristigasteridae Bleeker, 1872</p><p>TAXA INCLUDED AND RELATIONSHIPS. — ( Ilisha + Pellona) + ( Pristigaster + (( Chirocentrodon + Neoopisthopterus) + ( Pliosteostoma + ( Raconda + ( Opisthopterus + Odontognathus))))).</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Mouth superterminal (upward turned) (1:0&gt;2); supratemporal commissure passing through parietals not bony enclosed (7:1&gt;0); parietal-epioccipital ridge present (19:0&gt;1); posterior frontal fontanelles present (open) (27:0&gt;1); ethmopalatine cartilage present (46:0&gt;1); nasal small, bearing a bony flange to form a narrow groove for the supraorbital canal opening posteriorly (73:0&gt;1); sclerotic bones large, their total length representing almost the entire circumference of the orbit (82:0&gt;1); quadrate-mandibular articulation anterior to the orbit (131:0&gt;2); dorsal process of the ectopterygoid elongated and slender (135:0&gt;1); dorsal process of the posterior ceratohyal well-definite and large (147:0&gt;1); dorsal blade of the urohyal present and well developed (149:0&gt;1); ventral process of the second basibranchial well developed (178:0&gt;1); teeth on the first and second hypobranchials present (179:0&gt;1; 180:0&gt;1); basibranchial tooth plates 1-3 fused with the second basibranchial (182:0&gt;1); supraneurals oriented vertically or inclined forward (209:0&gt;1); dorsal fork of the epipleurals not projecting dorsally beyond the level of the epicentrals (225:1&gt;0); epicentrals and anterior ribs unfused (230:1&gt;0); pleural ribs/preural vertebrae ratio 0.20-0.40 or 0.41-0.46 (231:2&gt;0/1); interzygapophysal articulation absent (243:1&gt;0); postzygapophyses more developed than prezygapophyses (244:0&gt;2); anterior bony expansion in the parapophyses of the third vertebra present (245:0&gt;1); bony process on the first pleural rib present (248:0&gt;1); second and third postcleithra absent (263:1&gt;0; 264:1&gt;0); pelvic fin placed anterior to the dorsal-fin origin (280:1&gt;0); hypural diastema absent (288:0&gt;1); hypural 3 larger than hypural 1 (290:0&gt;1); prepelvic scutes anterior to the coracoid present (336:0&gt;1); 31-55 anal-fin pterygiophores (343:0&gt;1).</p><p>SUPPORT. — Relative Bremer = 78%.</p><p>REMARKS. — Nelson (1970) assigned the pristigasterids the rank of a superfamily due to their distinct and consistent gill arch characters (fusion of tooth plates with the second hypobranchials, and sometimes with the first, third, or both first and third hypobranchials, lacking other traits distinctive of engrauloids and clupeoids) which were supposed sufficiently unique to warrant a rank equivalent to other established clupeoid superfamilies, i.e., Chirocentroidea, Engrauloidea,and Clupeoidea .After the introduction of the superfamily “Pristigasteroidae” by Nelson (1970), Grande (1985) renamed the clade as Pristigasteroidea to standardize the suffix rather than reflect a fundamental change in the taxonomic composition of the group and added two new skeletal traits:supraneurals oriented either vertically or inclined anterodorsally and loss of the interlobar notch (hypural diastema) in the third hypural. Within the clade, Grande (1985) recognized three major groups at family level: 1) the Pristigasteridae (including “ Ilisha ” africana, Odontognathus, Opisthopterus, Pristigaster and Raconda) supported by the presence of a bony process on the first pleural rib which articulates with the shoulder girdle; and 2) the Pellonidae (including Chirocentrodon, Neoopisthopterus, Pellona and Pliosteostoma) supported by the maxillary-premaxillary gap covered by bone; and 3) the genus Ilisha (minus “ Ilisha ” africana). Although the monophyly of the Pristigasteroidea is well supported by several morphological and molecular based analyses (Sato 1994; Di Dario 2004b; Li &amp; Ortí 2007; Lavoué et al. 2013; Bloom &amp; Egan 2018; Egan et al. 2018; Wang et al. 2022) the relationships between the genera have not yet been comprehensively understood and these three groups have not been always recognized, mostly because pristigasteroids are usually undersampled in these analyses. Former parsimony-based phylogenetic analyses often recovered a clade formed by Chirocentrodon and Odontognathus in polytomy with Ilisha and Pellona (Sato 1994), or sister to Ilisha + Pellona (Vernygora 2020), or sister to Ilisha africana (Kevrekidis et al. 2024), or species of a paraphyletic Pellona that are successive sister of a clade formed by Ilisha and Opisthopterus (Di Dario 2004b) . Our tree topology about the intrarelationships of the monotypic Pristigasteroidea, however, is not comparable to most of the morphological or molecular analyses as these included only a few genera (mostly two to five), whereas our topology is, at least in part, consistent with that of Di Dario (1999). In conclusion, since the Pellonidae and Pristigasteridae sensu Grande (1985) have never been recognized in molecular or parsimony approaches, we confirm the monotypic nature of the Pristigasteroidea, including a single family ( Pristigasteridae) pending further detailed analyses. The putative fossil pristigasteroid † Gasteroclupea branisai Signeaux, 1964 has been re-described in detail by MarramÀ &amp; Carnevale (2017) who recovered this clupeomorph to be a member of the † Ellimmichthyiformes (see family Gasteroclupeidae), as confirmed also in this study.</p></div>	https://treatment.plazi.org/id/03E8879FB641FFDCFF3DF9B1FBF5F878	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB640FFDEFC5AF937FD06FB0E.text	03E8879FB640FFDEFC5AF937FD06FB0E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Clupeoidea Cuvier 1816	<div><p>Superfamily CLUPEOIDEA Cuvier, 1816, new usage</p><p>TAXA INCLUDED AND RELATIONSHIPS. — Hyperlophidae + ( Clupeidae + ( Ehiravidae + ( Alosidae + pan-Dorosomatidae))).</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Anterior arm of the orbitosphenoid long, reaching the lateral ethmoid (44:0&gt;1); posterior process of the lateral ethmoid present (65:0&gt;1); palatine teeth absent (114:1&gt;0); teeth or tooth plates on ceratobranchial 1 small (158:0&gt;1); teeth or tooth plates on ceratobranchials 2 and 3 small (160:0&gt;1); cartilage process of the anterior margin of the fourth ceratobranchial directed medially, forming a bridge over the ventral surface of the fourth basibranchial (161:0&gt;1); cartilage chevron at the distal tip of the epicentrals present (229:0&gt;1); parapophyses on third vertebra reduced (241:1&gt;0); medial process of the cleithrum present (258:0&gt;1); basal segments of the upper middle caudal-fin ray spatulate (317:0&gt;1); body scales of crenate type (361:0&gt;1); posterior margin of the body scales crenulate (362:0&gt;1).</p><p>SUPPORT. — Relative Bremer = 32%.</p><p>REMARKS. — The traditional composition of the superfamily Clupeoidea has been proposed by Grande (1985) to include the families Chirocentridae ( Chirocentrus) and his Clupeidae (Alosinae, Clupeinae, Dorosomatinae, Dussumieriinae, Pellonulinae) with the monophyly of the group thought to be supported by single condition, increase in pleural rib to preural vertebrae ratio (≥0.47), and possibly by a peculiar type of connection between the swim bladder and gut in a more posterior position of the esophagous with respect to other clupeiforms (Nelson 1970). The Clupeoidea sensu Grande (1985) was later proposed to be more closely related to the engraulids based on the presence of cartilage chevrons at the tips of the epicentrals, posteriorly directed parapophyses of the second vertebra, and interzygapophysal articulation (Di Dario 2002). Di Dario (2009) rejected the monophyly of the Clupeoidea sensu Grande (1985), in that Chirocentrus was supposed to be more closely related to the Engraulidae than to the other Clupeidae . Miyashita (2010) proposed an alternative hypothesis in which the Clupeoidea sensu Grande (1985) is closer to the Pristigasteroidea than to the Engrauloidea because they all share “a unique occipital articulation with the first vertebra”. Our analysis rejects the monophyly of the Clupeoidea and Clupeidae sensu Grande (1985), since Chirocentrus, Dussumieria, Etrumeus, Jenkinsia and Spratelloides are basal or more closely related to the Engraulidae than to the other clupeoids. For this reason, we propose a new usage for the superfamilial rank Clupeoidea that would include all the remaining clupeoid taxa.</p></div>	https://treatment.plazi.org/id/03E8879FB640FFDEFC5AF937FD06FB0E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB643FFDEFF0BFB16FAC2FECA.text	03E8879FB643FFDEFF0BFB16FAC2FECA.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Hyperlophidae Ogilby 1897	<div><p>Family Hyperlophidae Ogilby, 1897, new rank</p><p>TAXA INCLUDED AND RELATIONSHIPS. — († Gosiutichthys + † Knightia) + ( Hyperlophus + Potamalosa).</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — At least one predorsal scute present (318:0&gt;1); 11 to 20 predorsal scutes (326:3&gt;1).</p><p>SUPPORT. — Relative Bremer = 55%.</p><p>REMARKS. — The subfamily Hyperlophinae was established by Ogilby (1897) to contain Hyperlophus and Potamalosa, two small to medium sized double-armored herring genera inhabiting the marine and freshwater/brackish habitats of the Australia (Whitehead 1985). The subfamily has been diagnosed by the presence of complete abdominal and predorsal scute series and a single (posterior) supramaxilla, and a dorsal fin inserted above or just behind the pelvic-fin insertion (Ogilby 1897). In our analysis, the pair formed by the putative clupeine † Gosiutichthys and the putative pellonuline † Knightia from the Paleocene to Middle Eocene freshwater deposits of North America (Grande 1982b) is recovered sister to the extant Hyperlophinae, since most of them also share similar number of predorsal scutes (11-20). Despite recent molecular analyses recurrently recover the pair Hyperlophus + Potamalosa as sister to the clade formed by Clupea + Sprattus, in the family Clupeidae sensu Wang et al. (2022), our analysis might suggest that results from gene datasets might be biased by the lack of fossil data, since we recover these two pairs as successive sisters to the rest of clupeoids. The counterproof is that when fossil taxa are removed the pair ( Hyperlophus + Potamalosa) form a monophyletic group with the pair ( Clupea + Sprattus) (Fig. 41). For these reasons, we propose to keep the distinctiveness of these two pairs, elevating the subfamily Hyperlophinae Ogilby, 1897 at familial level, and considering Clupea and Sprattus as the sole members of the family Clupeidae Cuvier, 1816 .</p></div>	https://treatment.plazi.org/id/03E8879FB643FFDEFF0BFB16FAC2FECA	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB643FFDFFC45F8D0FE1AFD9C.text	03E8879FB643FFDFFC45F8D0FE1AFD9C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Clupeidae Cuvier 1816	<div><p>Family Clupeidae Cuvier, 1816, new usage</p><p>TAXA INCLUDED AND RELATIONSHIPS. — Clupea + Sprattus .</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Participation of the pterotic in the sub-epioccipital fossa (58:1&gt;0); anterior region of the mesethmoid keel perforated (62:0&gt;1); olfactory nerve foramen of the lateral ethmoid surrounded by the lateral ethmoid and mesethmoid (66:1&gt;2); teeth on the vomer present (69:0&gt;1); ventral process of the quadrate short (128:0&gt;1); foramen on the anterior ceratohyal produced by the overlap between lateral and medial foramina of the hyoid arterial duct present (140:0&gt;1); foramen in the fourth epibranchial present (163:0&gt;1); posterior concavity on the fourth epibranchial (166:0&gt;1); holes on the surface of the coracoid present (272:0&gt;1); keel of the pelvic scute absent (332:1&gt;0); more than 30 abdominal scutes (341:3&gt;4); focus of the body scales in postero-central position (360:2&gt;3).</p><p>SUPPORT. — Relative Bremer = 67%.</p></div>	https://treatment.plazi.org/id/03E8879FB643FFDFFC45F8D0FE1AFD9C	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB642FFDFFF36FBF8FC63FE21.text	03E8879FB642FFDFFF36FBF8FC63FE21.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Ehiravidae Deraniyagala 1929	<div><p>Family Ehiravidae Deraniyagala, 1929, new usage</p><p>TAXA INCLUDED AND RELATIONSHIPS. — Clupeonella + ( Ehiravinae + Pellonulinae).</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Skull roof unornamented (16:1&gt;0); posterior frontal fontanelles open and large (27:0&gt;1; 28:0&gt;1); teeth or tooth plates on ceratobranchial 1 absent (157:1&gt;0); teeth or tooth plates on ceratobranchials 2 and 3 absent (159:1&gt;0); medial process of the cleithrum absent (258:1&gt;0); caudal complex formed by fused PU1+U1 and autogenous U2 (295:0&gt;1); two epurals (304:0&gt;1); true cycloid body scales (361:1&gt;0); posterior margin of the body scales smooth (362:1&gt;0).</p><p>SUPPORT. — Relative Bremer = 56%</p><p>REMARKS. — The family Ehiravidae new usage comprises the subfamily Pellonulinae sensu Grande (1985) to the exclusion of Hyperlophus and Potamalosa but including Clupeonella (a clupeine according to Grande 1985). The Pellonulinae was the only subfamily of the Clupeidae, along with the Dussumieriinae, for which Grande (1985) provided a diagnosis, whereas the Alosinae, Dorosomatinae and Clupeinae were merely considered to be “groups of convenience” because of the difficulty to recognize unambiguous diagnostic characters. Grande (1985) diagnosed his Pellonulinae by the presence of a single ambiguous synapomorphy, the loss of the anterior supramaxilla, although he pointed out that “separation of this group based on this character is tenuous” because the anterior supramaxilla is absent also in other non-pellonuline clupeoids and because there are no other features that unite Potamalosa and Hyperlophus to the members of the pellonulines.Our analysis in agreement with many recent molecular analyses (Lavoué et al. 2013, 2014, 2017a; Bloom &amp; Lovejoy 2014; Wang et al. 2022) corroborates the molecular results, indicating that Potamalosa and Hyperlophus are not related to the Pellonulinae, whereas Clupeonella is resolved herein to be the basalmost ehiravid. However, contrary to molecular analyses that recovers the tribe Pellonulini sensu Grande (1985) nested within the Dorosomatidae, we recovered Clupeonella sister to the pair formed by the Ehiravidae sensu Wang et al. (2022) (here represented by Clupeichthys, Clupeonella, Corica and Gilchristella) and the Pellonulini sensu Grande (1985) (including Limnothrissa, Microthrissa, Nannothrissa, Odaxothrissa, Pellonula, Potamothrissa, and Stolothrissa) that are re-ranked herein at the subfamilial level (Fig. 44).</p></div>	https://treatment.plazi.org/id/03E8879FB642FFDFFF36FBF8FC63FE21	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB642FFDFFC96FC4DFB72FB7C.text	03E8879FB642FFDFFC96FC4DFB72FB7C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Ehiravinae Deraniyagala 1929	<div><p>Subfamily Ehiravinae Deraniyagala, 1929, new rank</p><p>TAXA INCLUDED AND RELATIONSHIPS. — Gilchristella + ( Clupeoides + ( Corica + Clupeichthys)).</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Lateral wings of the urohyal absent (151:1&gt;0); unique W-shaped articulation between the first and second vertebrae absent (207:1&gt;0); pelvic fins inserted anterior to the dorsal-fin origin (280:1&gt;0); lateral crest of hypural 1 absent (311:1&gt;0); 10-20 abdominal scutes (341:3&gt;2).</p><p>SUPPORT. — Relative Bremer = 57%.</p></div>	https://treatment.plazi.org/id/03E8879FB642FFDFFC96FC4DFB72FB7C	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB644FFD9FC7BFF15FB73FA56.text	03E8879FB644FFD9FC7BFF15FB73FA56.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Alosidae Svetovidov 1953	<div><p>Family Alosidae Svetovidov, 1953 (sensu Wang et al. 2022)</p><p>TAXA INCLUDED AND RELATIONSHIPS. — Sardina + ((† Eoalosa + Sardinops) + ( Alosa + Brevoortia)).</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Four discrete openings present in the otic region: the opening for the infraorbital sensory canal opens between sphenotic, pterotic and sometimes frontal; all other openings open on pterotic (32:7&gt;0); posterior pterotic spine present (57:0&gt;1); anterior border of the third infraorbital concave (79:0&gt;1); posterior cartilage of the fourth epibranchial formed by three cartilaginous elements fused into one with posterior process well developed, but neither anterior process nor medial expansion developed (167:3&gt;4); radial striations on the ventral part of the opercle (201:0&gt;1); more than 30 abdominal scutes (341:3&gt;4).</p><p>SUPPORT. — Relative Bremer = 56%.</p><p>REMARKS. — The Alosinae (shads) was one of the artificial subfamilies considered as “groups of convenience” by Grande (1985), since no osteological characters were recognized to support its monophyletic status. Traditionally, alosines included seven extant genera ( Alosa, Brevoortia, Ethmalosa, Ethmidium, Gudusia, Hilsa and Tenualosa) (Grande 1985; Whitehead 1985) united by a set of plesiomorphic characters, including their large size (up to 60 cm in SL), the possession of strong and well-developed abdominal scutes, and the presence of a median notch between the two contralateral premaxillae, plus some features of the gill arches and digestive tract (e.g., Nelson 1967a, b; Whitehead 1973, 1985). However, numerous molecular (Lavoué et al. 2007, 2014; Bloom &amp; Lovejoy 2014; Wang et al. 2022) and morphology-based analyses (Vernygora 2020; Kevrekidis et al. 2024) recurrently recovered Alosa and Brevoortia close to the putative clupeines Sardina and Sardinops, in agreement with our analyses that highlight seven unambiguous synapomorphies supporting this clade. Re-examination of the clupeomorph † Pugliaclupea nolardi from fossiliferous Upper Cretaceous (Campanian; Schlüter et al. 2008) levels of Nardò (southern Italy), considered as the oldest putative alosid or pan-alosid (Taverne 2004; Near &amp; Thacker 2024), ruled out this hypothesis since the holotype clearly shows diagnostic features of the † Ellimmichthyiformes, particularly of the † Armigatidae (G.M. pers. obs.).</p></div>	https://treatment.plazi.org/id/03E8879FB644FFD9FC7BFF15FB73FA56	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB644FFD9FF76FF15FE1AFDE7.text	03E8879FB644FFD9FF76FF15FE1AFDE7.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Pellonulinae Svetovidov 1952	<div><p>Subfamily Pellonulinae Svetovidov, 1952, new rank</p><p>TAXA INCLUDED AND RELATIONSHIPS. — Pellonula + ( Potamothrissa, Odaxothrissa, ( Stolothrissa + Limnothrissa), ( Microthrissa, Nannothrissa)).</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Ventral blade of the urohyal absent or inconspicuous (150:1&gt;0); posterior process of the fifth epibranchial absent or inconspicuous (168:1&gt;0); teeth or tooth plates over infrapharyngobranchial 4 absent (189:1&gt;0); distal region of most supraneurals greatly expanded (211:0&gt;1); articulation of the first rodlike postcleithrum (=second postcleithrum) with the supracleithrum well behind the cleithrum (268:0&gt;1).</p><p>SUPPORT. — Relative Bremer = 49%.</p></div>	https://treatment.plazi.org/id/03E8879FB644FFD9FF76FF15FE1AFDE7	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
03E8879FB647FFDAFC67FCB7FB72FB57.text	03E8879FB647FFDAFC67FCB7FB72FB57.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dorosomatidae Bleeker 1872	<div><p>Family Dorosomatidae Bleeker, 1872, new usage</p><p>TAXA INCLUDED AND RELATIONSHIPS. — ( Escualosa + († Chasmoclupea + Rhinosardinia)) + (( Platanichthys + Lile) + (( Amblygaster + ( Harengula + Herklotsichthys)) + ( Opisthonema + ( Sardinella + ( Hilsa + ( Tenualosa + ( Dorosoma + ( Konosirus + ( Clupanodon + ( Anodontostoma + Nematalosa)))))))))).</p><p>UNAMBIGUOUS SYNAPOMORPHIES. — Distal region of the supraneurals greatly expanded (211:0&gt;1); membranous anterodorsal outgrowth of uroneural 1 absent (299:2&gt;0).</p><p>SUPPORT. — Relative Bremer = 70%.</p></div>	https://treatment.plazi.org/id/03E8879FB647FFDAFC67FCB7FB72FB57	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Marramà, Giuseppe;Carnevale, Giorgio	Marramà, Giuseppe, Carnevale, Giorgio (2026): A phylogeny and revised classification of clupeomorph fishes (Teleostei, Otomorpha). Zoosystema 48 (17): 401-556, DOI: 10.5252/zoosystema2026v48a17
