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![Caudal skeletons of fossil elopiforms. A, †Anaethalion knorri (JME SOS 2282; Upper Jurassic), figure reversed to the left. B, †Elopsomolos sp. (NHM 37048; Upper Jurassic). Scales = 2 mm. Abbreviations: d, hypural diastema; dsc, dorsal scute; E1-3, epurals of ural centra 1 P , 2 P and 3 P ; ff, fringing fulcra; H1-3, hyp urals 1-3; hy, hypuraphophysis; naPU1, U1, neural arches of preural 1 and ural 1 P ; naU1-2, neural arch of ural centrum 1+2 P ; nsPU2, neural spine of preural centrum 2; PH, parhypural or haemal spine of preural centrum 1; 1PR, first principal ray; PU6-1, preural centra 6-1; U1+2, ural centrum 1+2 P ; U1+2+H1, 2, ural centrum 1 P +2 P +hypurals 1 and 2; UN1-4 [UN4-7], uroneurals 1-4 (position) → [modified neural arches of ural centra 4 to 7 P ]; vscu, ventral caudal scute.](profile/Gloria-Arratia/publication/259496648/figure/fig4/AS:669391872069644@1536606867449/Caudal-skeletons-of-fossil-elopiforms-A-Anaethalion-knorri-JME-SOS-2282-Upper.png)
Caudal skeletons of fossil elopiforms. A, †Anaethalion knorri (JME SOS 2282; Upper Jurassic), figure reversed to the left. B, †Elopsomolos sp. (NHM 37048; Upper Jurassic). Scales = 2 mm. Abbreviations: d, hypural diastema; dsc, dorsal scute; E1-3, epurals of ural centra 1 P , 2 P and 3 P ; ff, fringing fulcra; H1-3, hyp urals 1-3; hy, hypuraphophysis; naPU1, U1, neural arches of preural 1 and ural 1 P ; naU1-2, neural arch of ural centrum 1+2 P ; nsPU2, neural spine of preural centrum 2; PH, parhypural or haemal spine of preural centrum 1; 1PR, first principal ray; PU6-1, preural centra 6-1; U1+2, ural centrum 1+2 P ; U1+2+H1, 2, ural centrum 1 P +2 P +hypurals 1 and 2; UN1-4 [UN4-7], uroneurals 1-4 (position) → [modified neural arches of ural centra 4 to 7 P ]; vscu, ventral caudal scute.
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Citations
... Terminology for the elements of the caudal skeleton follows Arratia (2008) and Schultze & Arratia (2013). Meristic data follow Fink & Weitzman (1974). ...
... Opsanus tau (Linnaeus, 1766) (Batrachoidinae; Fig. 2) is used to illustrate the general morphology of the caudal skeleton of Batrachoidiformes. Terminology for caudal-fin rays and skeletal elements in general follows that of Arratia (2008) and Schultze & Arratia (2013), with the exception of the numbering of hypurals, which we number sequentially rather than presuppose hypotheses of homology to other groups of teleosts (see Hilton et al., 2015). ...
... The hypurapophysis is a process that projects laterally from the arches of the parhypural (Nursall, 1963;Schultze & Arratia, 2013) and serves as an attachment site for the m. hypochordal longitudinalis (Winterbottom, 1974). ...
The caudal-fin skeleton is a primary data source for systematics of fishes, with characters from this complex being proposed as synapomorphies at many taxonomic levels. Batrachoidiformes is recognized as monophyletic, although intraordinal relationships are unclear. Likewise, interrelationships of Batrachoidiformes to other percomorphs are not well established. The caudal skeleton of Batrachoidiformes has not been thoroughly studied and is poorly represented in recent phylogenetic analyses. In this study, we examined the caudal-fin skeleton of 55 of the 82 species and 22 of the 23 genera of Batrachoidiformes, emphasizing the detection of intraspecific variation to recognize morphological characters with phylogenetic significance. Intraspecific variation is high, especially in the shape of epurals and the parhypural flange. A dorsal prezygapophysis on the first ural centrum and the acute articular edge of the parhypural flange are interpreted as putative synapomorphies of Porichthyinae. The anterior epural supporting two procurrent fin rays is found only in some Halophryninae, but is absent in Allenbatrachus, Batrachomeus, Batrichthys and Halophryne. Among Batrachoidiformes, a hypurapophysis-like process on the first ural centrum is found in Thalassophryninae and Barchatus, Batrichthys, Bifax, Chatrabus, Colletteichthys, Halobatrachus, Perulibatrachus and Riekertia. Caudal-fin ray counts are phylogenetically informative at several taxonomic levels. Distal caudal cartilages are described for Batrachoidiformes for the first time.
... Thus, terminology based on homologies of skull-roof bones are not followed here. The diural terminology of the caudal endoskeleton is followed instead of the polyural form; the numbering of uroneurals does not imply homology with other teleostean clades (see Schultze & Arratia 2013, Wiley et al. 2015, only position. The terminology of the vertebral column and associated structures, and of the fin rays, follow Arratia (1997Arratia ( , 2008 and . ...
... Principal rays 5 and 6 are supported by hypural 5; hypural 4 supports principal rays 7 to 9; principal ray 10 is supported by hypural 3; principal ray 11 is also associated with hypural 3; hypural 2 supports principal rays 12 and 13; hypural 1 supports principal rays 14 to 17; the parhypural supports principal rays 18 and 19. There is a gap between the base of the first procurrent ray and the first principal ray, which is associated with the trajectory of the notochord (P30920a) (Schultze & Arratia 2013). The bases of principal rays in the centre of the fin, i.e., principal rays 9, 10 and 11, are enlarged, but they do not have dorsal processes. ...
Bean, L. B. & Arratia, G., 4 October 2019. Anatomical revision of the Australian teleosts Cavenderichthys talbragarensis and Waldmanichthys koonwarri impacting on previous phylogenetic interpretations of teleostean relationships. Alcheringa 44, 121-159. DOI: 10.1080/03115518.2019.1666921.
Australia has two important sites for Mesozoic fishes. The Talbragar Fossil Fish Bed in New South Wales is a Tithonian freshwater lake deposit containing the iconic form Cavenderichthys talbragarensis, first described in 1895. The Koonwarra Fossil Bed in Victoria is an Albian freshwater lake deposit containing Waldmanichthys koonwarri, first described in 1971. Following the tradition of the time, both species were first ascribed to the historic genus Leptolepis. In 2015, these two species were placed into a newly erected family, Luisiellidae, with the Oxfordian–Tithonian fish Luisiella feruglioi from the Cañadón Calcáreo Formation in Chubut, Argentinian Patagonia. This new family was interpreted as a stem teleost group, closer to Leptolepis coryphaenoides than to the crown-group Teleostei, a contrary hypothesis to previous interpretations that placed Cavenderichthys as a teleost incertae sedis in the crown Teleostei. This paper re-examines the morphological characters of the Australian taxa, including 46 previously undescribed specimens of W. koonwarri, from Museum Victoria. Some of the new characters include the special configuration of the jaws and the position of the quadrate-mandibular articulation; the special vertebral pattern at the level of the abdominal/caudal regions; a stegural-like uroneural in the caudal skeleton; and the structure of the scales. Finding the new characters in Waldmanichthys called for reappraisals of the morphology of Cavenderichthys and Luisiella. In the case of Cavenderichthys, 34 specimens were re-examined, but for specimens of Luisiella, a conservative approach was followed, based on its last morphological description as well as photographs taken more recently. The systematic position of the three Gondwanan taxa was re-evaluated using a pre-existing data matrix including 240 characters and 56 taxa. The new results give a very different scenario, with the three taxa now included in the crown-group Teleostei. The family Luisiellidae is restricted to its type species L. feruglioi. The two Australian fish genera cluster together with the Late Jurassic European genera Leptolepides and Orthogonikleithrus and are now ascribed to the family Orthogonikleithridae. The new results suggest that the three Gondwanan genera are stem taxa to the Osteoglossocephala (osteoglosomorphs plus more advanced teleosts), and their combination of morphological characters has a major effect on the interpretation of basal euteleosts, questioning some previous interpretations, as for instance, the homology of the stegural as an euteleostean character.
Lynne B. Bean* [lynne.bean@anu.edu.au], Research School of Earth Sciences, Australian National University, Acton 2001, Australia; Gloria Arratia [garratia@ku.edu], Department of Ecology and Evolutionary Biology and Biodiversity Institute, University of Kansas, Dyche Hall, Lawrence, KS 66045, USA.
... 18, 19A-D), and in ostariophysans (e.g., Monod, 1968;Lundberg, Baskin, 1969;Fink, Fink, 1981;Schultze, Arratia, 1989, 2013Fujita, 1990). However, it is unknown whether the structure interpreted as a compound centrum in osta- riophysans forms the same way in different ostariophysan subgroups (see Schultze, Arratia, 2013;Wiley et al., 2015). The fossil ostariophysan †Tischlingerichthys also has a long vertebral centrum bearing the parhypural and hypurals 1 and 2 (Arratia, 1997: fig. ...
A morphological revision is presented here on the cohort Otomorpha, a clade currently interpreted as the most primitive among the large supercohort Clupeocephala. Otomorpha is a morphologically heterogeneous group represented by clupei forms , alepocephaliforms, and ostariophysans (gonorynchiforms, cypriniforms, characiforms, siluriforms, and gymnoti forms) that inhabit various marine and freshwater environments worldwide. Otomorphs have a long (ca. 145 Ma) and diverse fossil record. They are the largest fish teleostean clade worldwide, as well as the largest of the Neotropical Region. While molecular studies strongly confirm the monophyly of Otomorpha, most potential morphological synapomorphies of the group become homoplastic largely due to the peculiar morphological character states (either losses or transformations) present in alepocephaliforms. The fusion of haemal arches with their respective vertebral centra anterior to preural centrum 2 stands as an unambiguous synapomorphy of the clade. The ankylosis or fusion of the extrascapular and parietal bones, and silvery areas associated with the gas bladder are also interpreted as synapomorphies, although they are homoplastic characters mainly due to secondary losses or further transformations of the morphological features in the alepocephaliforms.
... Here, we analyzed, in a phylogenetic context, newly described caudal fin skeletal features among gars (Lepisosteiformes) (Desvignes et al., 2018) and their potential role in the establishment of symmetry in neopterygians and homocercality in teleosts. More specifically, we focused attention on previously unde- scribed endoskeletal and mesodermal features associated with hypurals 2 and 3, recalling the hypural diastema in teleosts (Arratia, 2013;Schultze and Arratia, 2013) by including a consis- tent gap between two adjacent hypurals that separates a pair of plates of connective tissue, and dermoskeletal features like the earliest-forming pair of caudal lepidotrichia, which we find to develop at hypurals 1 and 2 in contrast to previous description at hypurals 2 and 3 (Metscher and Ahlberg, 2001;Desvignes et al., 2018). ...
... During development and in adult specimens, hypurals were generally evenly spaced and they decreased in size progressively from anterior to posterior. We observed, however, that the space between hypurals 2 and 3 was generally greater than between other neighboring hypurals, which was more evident in juveniles, and resembled the hypural diastema described in teleosts (Arratia, 2013;Schultze and Arratia, 2013). The size and shape difference between hypurals 2 and 3 was also greater than the difference between any other pair of adjacent hypurals (Figs. ...
... Between 9 and 10 dpf (approximately 5 mm TL), the proximal epiphyses of hypural 1 and the parhypural fused, and the first ural centrum began to mineralize as a compound centrum of preural centrum 1 and ural centra 1 and 2. At approximately the same age, the opisthural cartilage and the epurals appeared and the central lepidotrichia began to mineralize, forming anterior and posterior to the separation between hypural 2 and 3 ( Fig. 2E- G) which is known as the hypural diastema (Arratia, 2013;Schultze and Arratia, 2013). Ural centrum 1 mineralized well before the anterior-to-posterior sequence of mineralizing caudal verte- brae had reached preural centrum 1 (Fig. 2F,G), in agreement with previous observations (Bird and Mabee, 2003;BensimonBrito et al., 2012a,b). ...
Background:
The caudal fin of actinopterygians transitioned from a heterocercal dorsoventrally asymmetrical fin to a homocercal externally symmetrical fin in teleosts through poorly understood evolutionary developmental mechanisms. We studied the caudal skeleton of major living actinopterygian lineages, including polypteriformes, acipenseriformes, Holostei (gars and bowfin), and teleosts, compared to reports of extinct neopterygians and basal teleosteans. We focused on the hypural diastema complex, which includes 1) a gap between hypurals 2 and 3, that 2) separates two plates of connective tissue at 3) the branching of caudal vasculature; these features had been considered as a shared, derived trait of teleosts, a synapomorphy.
Results:
These studies revealed that gars and teleosts share all three features of the hypural diastema complex. Absence of a complex with these features from bowfin, fossil Holostei, and stem Teleostei argues in favor of repetitive, independent emergence in several neopterygian and basal Teleostei lineages, or less likely, many independent losses. We further observed that in gars and teleosts, the earliest developing lepidotrichia align with the horizontal adult body axis, thus participating in external symmetry.
Conclusions:
These results suggest that the hypural diastema complex in teleosts and gars represents a homoplasy among neopterygians and that it emerged repeatedly by parallel evolution due to shared inherited underlying genetic and developmental programs (latent homology). Because the hypural diastema complex exists in gars with heterocercal tails, this complex is independent of homocercality. This article is protected by copyright. All rights reserved.
... Evolution of the dorsal and anal fins occurred con- comitantly with other morphological innovations, in the skull and mouth for example, which provided improvements in feeding capacities, leading to the rise of the neopterygians as the taxo- nomically dominant group of vertebrates (L opez-Arbarello, 2012). Among neopterygians, the largely dominant Teleost infra- class represents more than 99.9% of living actinopterygian spe- cies (Eschmeyer, 2015;Nelson, 2006).Teleosts are characterized morphologically by numerous lineage-specific characters, including additional innovations in the jaw that further improved feeding, as well as the emergence of a homocercal caudal fin (Arratia, 2015;Sallan, 2014;Schultze and Arratia, 1989;Schultze and Arratia, 2013), which may have contributed to the successful radiation of teleosts into almost all aquatic ecosystems. Indeed, the caudal fin is a major component of mobility in fish because it generates swimming power and contributes to maneuverability, thus helping to control swimming speed and direction of move- ment, which facilitate prey capture and predator avoidance (Flammang and Lauder, 2009;Lauder, 2000). ...
... Indeed, the caudal fin is a major component of mobility in fish because it generates swimming power and contributes to maneuverability, thus helping to control swimming speed and direction of move- ment, which facilitate prey capture and predator avoidance (Flammang and Lauder, 2009;Lauder, 2000). The caudal fin and its supporting internal skeleton are highly variable among acti- nopterygians, so different skeletal elements, their morphological variations, and their associations with each other are frequently used in a variety of ecological, biomechanical, and systematic studies (e.g., Arratia and Schultze, 1992;Cloutier et al., 2011;Cloutier and Arratia, 2004;Fiaz et al., 2012;Flammang and Lauder, 2009;Gr€ unbaum et al. 2003;Lauder, 2000;Schultze and Arratia, 1986;Schultze and Arratia, 1988;Schultze and Arratia, 2013). ...
... Despite numerous studies on caudal fin skeletons (e.g., Metscher and Ahlberg, 2001;Moriyama and Takeda, 2013;Schultze and Arratia, 2013), the morphological and genetic pro- cesses by which the caudal fin transitioned from the heterocercal state in basal actinopterygians to the homocercal form in teleosts remains not well understood. This information gap is due largely to several factors. ...
Background:
The caudal fin of actinopterygians experienced substantial morphological changes during evolution. In basal actinopterygians, the caudal fin skeleton supports an asymmetrical heterocercal caudal fin, while most teleosts have a symmetrical homocercal caudal fin. The transition from the ancestral heterocercal form to the derived homocercal caudal fin remains poorly understood. Few developmental studies provide an understanding of derived and ancestral characters among basal actinopterygians. To fill this gap, we examined the development of the caudal fin of spotted gar Lepisosteus oculatus, one of only eight living species of Holostei, the sister group to the teleosts.
Results:
Our observations of animals from fertilization to more than a year old provide the most detailed description of the developmentof caudal fin skeletal elements in any Holostean species. We observed two different types of distal caudal radials replacing two transient plates of connective tissue, identifying two hypaxial ensembles separated by a space between hypurals 2 and 3. These features have not been described in any gar species, but can be observed in other gar species, and thus represent anatomical structures common to lepisosteiformes.
Conclusion:
The present work highlights the power and importance of ontogenic studies, and provides bases for future evolutionary and morphological investigations on actinopterygians fins. This article is protected by copyright. All rights reserved.
... The terms of the fins and associated bones is used after Arratia (2008Arratia ( , 2009). The terminology of the caudal skeleton and count of vertebrae follows Nybelin (1963), Arratia and Schultze (1992) and Schultze and Arratia (2013). The classification of scales follows Schultze (2016). ...
... Cephalic sensory canals were studied following Northcutt (1989). The postcranial anatomy (vertebral column) was described following Arratia et al. (2001); clavicle and serrated appendage follow Arratia (2013); caudal skeleton terminology follows Schultze and Arratia (2013); fin rays and fulcra follows Arratia (2008); and that of scales follows Schultze (1966;. The fishes studied here present some enlarged, thin scales that are positioned anteromedian and posteromedian to the dorsal fin. ...
Although the Cretaceous is characterized by a rich fish diversity, Cretaceous continental fishes from Gondwana are poorly known and comparatively scarce. Among these fishes, the family Pleuropholidae is only known by a few species relatively poorly preserved, from the Middle Jurassic to Lower Cretaceous of Europe, Africa, North America, and South America. In this paper, two new species of the pleuropholid new genus Zurupleuropholis are described, Z. quijadensis gen. et sp. nov. and Z. decollavi gen. et sp. nov. The new fishes were recovered in the Lower Cretaceous lacustrine Lagarcito Formation of central-west Argentina. This taxon constitutes a relevant finding considering that the representation of the family Pleuropholidae is rare worldwide. Zurupleuropholis gen. nov. appears to be the youngest known member of Pleuropholidae, and it represents the second record of the family in South America and the first record in the Cretaceous of the continent.
... Structures of the vertebral centrum and the vertebral centrum-associated elements are defined in Table 1, following Schultze & Arratia (2013) and other authors listed in the table. Cartilages of the vertebral centra and associated elements were defined following the classification of Witten et al. (2010). ...
Teleost vertebral centra are often similar in size and shape, but vertebral-associated elements, i.e. neural arches, haemal arches and ribs, show regional differences. Here we examine how the presence, absence and specific anatomical and histological characters of vertebral centra-associated elements can be used to define vertebral column regions in juvenile Chinook salmon (Oncorhynchus tshawytscha). To investigate if the presence of regions within the vertebral column is independent of temperature, animals raised at 8 and 12 °C were studied at 1400 and 1530 degreedays, in the freshwater phase of the life cycle. Anatomy and composition of the skeletal tissues of the vertebral column were analysed using Alizarin red S whole-mount staining and histological sections. Six regions, termed I–VI, are recognised in the vertebral column of specimens of both temperature groups. Postcranial vertebrae (region I) carry neural arches and parapophyses but lack ribs. Abdominal vertebrae (region II) carry neural arches and ribs that articulate with parapophyses. Elastic- and fibrohyaline cartilage and Sharpey's fibres connect the bone of the parapophyses to the bone of the ribs. In the transitional region (III) vertebrae carry neural arches and parapophyses change stepwise into haemal arches. Ribs decrease in size, anterior to posterior. Vestigial ribs remain attached to the haemal arches with Sharpey's fibres. Caudal vertebrae (region IV) carry neural and haemal arches and spines. Basidorsals and basiventrals are small and surrounded by cancellous bone. Preural vertebrae (region V) carry neural and haemal arches with modified neural and haemal spines to support the caudal fin. Ural vertebrae (region VI) carry hypurals and epurals that represent modified haemal and neural arches and spines, respectively. The postcranial and transitional vertebrae and their respective characters are usually recognised, but should be considered as regions within the vertebral column of teleosts because of their distinctive morphological characters. While the number of vertebrae within each region can vary, each of the six regions is recognised in specimens of both temperature groups. This refined identification of regionalisation in the vertebral column of Chinook salmon can help to address evolutionary developmental and functional questions, and to support applied research into this farmed species.
... Counst of the holotype are given in parentheses. Meristic and morphometric data, nomenclature in the osteology, myology and cephalic laterosensory system follows the most recent contributions in that sense [13,14,15,16]. The new taxon plus the putative close species T. crassicaudatus, T. igobi, and T. stawiarski are treated as Trichomycterus stawiarski group in the text. ...
A new species assigned to the genus Trichomycterus from the area of the waterfalls of
Tabay stream, Parana´ River basin, Misiones, Argentina, is described. Trichomycterus ytororo sp. nov. is distinguished from all other species in the genus by the presence of 31–35 dorsal procurrent caudal-fin rays and the combination of some external characters such as: coloration, number of pectoral–fin rays and pores of the laterosensory canals. The new taxon belongs to a presumably monophyletic group of species composed of T. crassicaudatus, T. igobi, and T. stawiarski based on the presence of 24 or more thickly ossified and rigid procurrent caudal-fin rays with a slender distal tip extending along the tips of at least ten neural spines.
... Cleithral bulge and posterior cleithral process weakly developed on day 6, and well developed by the end of postflexion stage. Spurious ray [20] present at the end of pectoral-fin spine since early in larval development (9 days), and persist in adults. Pelvic-fin bud rounded, base large at the start of postflexion stage (day 4). ...
Pseudopimelodidae are Neotropical catfishes characterized by having slightly to strongly depressed body in fully developed specimens. The largest species of the family with 500 mm SL, Lophiosilurus alexandri, experiences impressive changes in body shape during development, becoming extremely depressed when fully developed. Accordingly, Lophiosilurus alexandri is an ideal species to observe the morphological changes during ontogeny, and to seek solid interpretations on the polarity of characters. Specimens of distinct larval periods (yolk sac, flexion and postflexion; n = 186 specimens) and juvenile stages (n = 20) were analyzed. Changes in body shape, position of mouth and eye, morphology of fins and pigmentation were observed during the development of Lophiosilurus. Larvae (5.7-11.2 mm standard length) had pigmentation concentrated on the head and parts of body, eyes small and pigmented, short barbels, and well-developed finfold. Juveniles (15.9-28.1 mm standard length) had body shape similar to adult, with head depressed and bearing bony ridges, large mouth, dorsally-oriented eyes, small barbels and well-developed shoulder bulges (cleithral width). The greatest morphological changes in the development of L. alexandri occurred during the postflexion larval stage. Relative to standard length, measurements of snout length, head depth and body depth are smaller in juveniles than in larvae, but body width is larger. New interpretations on the phylogenetic characters related to these changes are provided in view of the two alternative hypotheses of the evolution of Pseudopimelodidae.
