Morphological Features of Cephalic Skeleton of the Adult Black Sea Turbot (Kalkan) Scophthalmus maximus var. maeotica (Pleuronectiformes, Scophthalmidae)

For the first time the full description of neural and visceral cephalic skeleton of adult Black Sea turbot (kalkan), Scophthalmus maximus (Pallas) has been carried out. The detailed outline of the norm of development of cephalic skeleton in adult Black Sea turbot with up-to-date nomenclature of bone...

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Datum:	2013
Hauptverfasser:	Yelnikov, D.V., Khanaychenko, A.N.
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Sprache:	English
Veröffentlicht:	Інститут зоології ім. І.І. Шмальгаузена НАН України 2013
Schriftenreihe:	Вестник зоологии
Schlagworte:	Морфология
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Zitieren:	Morphological Features of Cephalic Skeleton of the Adult Black Sea Turbot (Kalkan) Scophthalmus maximus var. maeotica (Pleuronectiformes, Scophthalmidae) / D.V. Yelnikov, A.N. Khanaychenko // Вестник зоологии. — 2013. — Т. 47, № 5. — С. 441–450. — Бібліогр.: 7 назв. — англ.

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spelling	irk-123456789-1098382016-12-18T03:02:40Z Morphological Features of Cephalic Skeleton of the Adult Black Sea Turbot (Kalkan) Scophthalmus maximus var. maeotica (Pleuronectiformes, Scophthalmidae) Yelnikov, D.V. Khanaychenko, A.N. Морфология For the first time the full description of neural and visceral cephalic skeleton of adult Black Sea turbot (kalkan), Scophthalmus maximus (Pallas) has been carried out. The detailed outline of the norm of development of cephalic skeleton in adult Black Sea turbot with up-to-date nomenclature of bone elements is offered as a basis to conduct further studies on variability of skeleton elements and abnormalities among the Black Sea turbot morphotypes from natural populations and artificially reared specimens. Впервые представлено полное описание нормы строения черепной коробки и висцерального скелета взрослых особей черноморской камбалы калкан, Scophthalmus maximus (Pallas). С учётом современной номенклатуры составлена подробная схема стандартного строения головного отдела скелета калкана, которую в дальнейшем предлагается использовать как основу нормы строения скелета при изучении изменчивости морфотипов и аномалий скелета особей черноморского калкана из природных популяций и выращенных в искусственных условиях. 2013 Article Morphological Features of Cephalic Skeleton of the Adult Black Sea Turbot (Kalkan) Scophthalmus maximus var. maeotica (Pleuronectiformes, Scophthalmidae) / D.V. Yelnikov, A.N. Khanaychenko // Вестник зоологии. — 2013. — Т. 47, № 5. — С. 441–450. — Бібліогр.: 7 назв. — англ. 0084-5604 DOI 10.2478/vzoo-2013-0047 http://dspace.nbuv.gov.ua/handle/123456789/109838 597.556.35:591.4(262.5) en Вестник зоологии Інститут зоології ім. І.І. Шмальгаузена НАН України
institution	Digital Library of Periodicals of National Academy of Sciences of Ukraine
collection	DSpace DC
language	English
topic	Морфология Морфология
spellingShingle	Морфология Морфология Yelnikov, D.V. Khanaychenko, A.N. Morphological Features of Cephalic Skeleton of the Adult Black Sea Turbot (Kalkan) Scophthalmus maximus var. maeotica (Pleuronectiformes, Scophthalmidae) Вестник зоологии
description	For the first time the full description of neural and visceral cephalic skeleton of adult Black Sea turbot (kalkan), Scophthalmus maximus (Pallas) has been carried out. The detailed outline of the norm of development of cephalic skeleton in adult Black Sea turbot with up-to-date nomenclature of bone elements is offered as a basis to conduct further studies on variability of skeleton elements and abnormalities among the Black Sea turbot morphotypes from natural populations and artificially reared specimens.
format	Article
author	Yelnikov, D.V. Khanaychenko, A.N.
author_facet	Yelnikov, D.V. Khanaychenko, A.N.
author_sort	Yelnikov, D.V.
title	Morphological Features of Cephalic Skeleton of the Adult Black Sea Turbot (Kalkan) Scophthalmus maximus var. maeotica (Pleuronectiformes, Scophthalmidae)
title_short	Morphological Features of Cephalic Skeleton of the Adult Black Sea Turbot (Kalkan) Scophthalmus maximus var. maeotica (Pleuronectiformes, Scophthalmidae)
title_full	Morphological Features of Cephalic Skeleton of the Adult Black Sea Turbot (Kalkan) Scophthalmus maximus var. maeotica (Pleuronectiformes, Scophthalmidae)
title_fullStr	Morphological Features of Cephalic Skeleton of the Adult Black Sea Turbot (Kalkan) Scophthalmus maximus var. maeotica (Pleuronectiformes, Scophthalmidae)
title_full_unstemmed	Morphological Features of Cephalic Skeleton of the Adult Black Sea Turbot (Kalkan) Scophthalmus maximus var. maeotica (Pleuronectiformes, Scophthalmidae)
title_sort	morphological features of cephalic skeleton of the adult black sea turbot (kalkan) scophthalmus maximus var. maeotica (pleuronectiformes, scophthalmidae)
publisher	Інститут зоології ім. І.І. Шмальгаузена НАН України
publishDate	2013
topic_facet	Морфология
url	http://dspace.nbuv.gov.ua/handle/123456789/109838
citation_txt	Morphological Features of Cephalic Skeleton of the Adult Black Sea Turbot (Kalkan) Scophthalmus maximus var. maeotica (Pleuronectiformes, Scophthalmidae) / D.V. Yelnikov, A.N. Khanaychenko // Вестник зоологии. — 2013. — Т. 47, № 5. — С. 441–450. — Бібліогр.: 7 назв. — англ.
series	Вестник зоологии
work_keys_str_mv	AT yelnikovdv morphologicalfeaturesofcephalicskeletonoftheadultblackseaturbotkalkanscophthalmusmaximusvarmaeoticapleuronectiformesscophthalmidae AT khanaychenkoan morphologicalfeaturesofcephalicskeletonoftheadultblackseaturbotkalkanscophthalmusmaximusvarmaeoticapleuronectiformesscophthalmidae
first_indexed	2025-07-07T23:43:28Z
last_indexed	2025-07-07T23:43:28Z
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fulltext	UDC 597.556.35:591.4(262.5) MORPHOLOGICAL FEATURES OF CEPHALIC SKELETON OF THE ADULT BLACK SEA TURBOT (KALKAN) SCOPHTHALMATUS MAXIMUS VAR. MAEOTICA (PLEURONECTIFORMES, SCOPHTHALMIDAE) D. V. Yelnikov, A. N. Khanaychenko Institute of Biology of Southern Seas A. O. Kovalevsky, NAS of Ukraine, Nakhimov pr., 2, Sevastopol, 99011 Ukraine E-mail: a.khanaychenko@gmail.com Morphological Features of Cephalic Skeleton of the Adult Black Sea Turbot (Kalkan) Scophthalmus max- imus var. maeotica (Pleuronectiformes, Scophthalmidae). Yelnikov D. V., Khanaychenko A. N. — For the first time the full description of neural and visceral cephalic skeleton of adult Black Sea turbot (kalkan), Scophthalmus maximus (Pallas) has been carried out. The detailed outline of the norm of development of cephalic skeleton in adult Black Sea turbot with up-to-date nomenclature of bone elements is offered as a basis to conduct further studies on variability of skeleton elements and abnormalities among the Black Sea turbot morphotypes from natural populations and artificially reared specimens. Key wo rd s: the Black Sea turbot, morphological features, cephalic skeleton. Ìîðôîëîãè÷åñêèå îñîáåííîñòè ñòðîåíèÿ ãîëîâíîãî îòäåëà ñêåëåòà ÷åðíîìîðñêîé êàìáàëû êàëêàí, Scophthalmus maximus var. maeotica (Pleuronectiformes, Scophthalmidae). Åëüíèêîâ Ä. Â., Õàíàé÷åíêî À. Í. — Âïåðâûå ïðåäñòàâëåíî ïîëíîå îïèñàíèå íîðìû ñòðîåíèÿ ÷åðåïíîé êîðîáêè è âèñöåðàëüíîãî ñêåëåòà âçðîñëûõ îñîáåé ÷åðíîìîðñêîé êàìáàëû êàëêàí, Scophthalmus maximus (Pallas). Ñ ó÷¸òîì ñîâðåìåííîé íîìåíêëàòóðû ñîñòàâëåíà ïîäðîáíàÿ ñõåìà ñòàíäàðòíîãî ñòðîåíèÿ ãîëîâíîãî îòäåëà ñêåëåòà êàëêàíà, êîòîðóþ â äàëüíåéøåì ïðåäëàãàåòñÿ èñïîëüçîâàòü êàê îñíîâó íîðìû ñòðîåíèÿ ñêåëåòà ïðè èçó÷åíèè èçìåí÷èâîñòè ìîðôîòèïîâ è àíîìàëèé ñêåëåòà îñîáåé ÷åðíîìîðñêîãî êàëêàíà èç ïðèðîäíûõ ïîïóëÿöèé è âûðàùåííûõ â èñêóññòâåííûõ óñëîâèÿõ. Êëþ÷åâûå ñëîâà: ÷åðíîìîðñêàÿ êàìáàëà êàëêàí, ìîðôîëîãè÷åñêèå îñîáåííîñòè, ñêåëåò ãîëîâû. Introduction Morphological peculiarities of fish skeleton should be undoubtedly considered while carrying out analysis of intraspecific variability of the species within its natural habitat, and for analysis of deformities induced dur- ing early ontogenesis under adverse environmental parameters in natural, or in artificial conditions. Studies of morphological variability in Scophthalmus maximus (syn. Psetta maximus) var. maeotica (Pallas, 1814) (Pleu- ronectiformes, Scophthalmidae) (kalkan, or Black Sea turbot, further referred to as BST) carried out in the Institute of Biology of Southern Seas, Sevastopol, Ukraine since 2006 revealed various abnormalities in mor- phology (including skeleton) of BST from natural population (Khanaychenko et al., 2008), and comparative study of abnormalities in the wild and artificially reared specimens in order to reduce skeleton deformities in larviculture reared BST juveniles, was carried out (Khanaychenko et al., in prep.). In the only publication concerning morphology of BST skeleton (Kalinina, 1959), the incomplete description of the cranial part of cephalic skeleton of BST was presented that was considered insufficient for analysis of intraspecific variability and skeleton pathologies of this species. Both trends of morphological research require standardized approach to analysis of skeleton morphology. Detailed description of the normal structure of cephalic skeleton of BST according to up-to-date nomenclature should contribute to better further analysis of the levels of morphological variability and pathology of these species both from nature and aquaculture. Vestnik zoologii, 47(5): e-42–e-51, 2013 DOI 10.2478/vzoo-2013-0047 Unauthenticated Download Date \| 12/15/16 7:15 PM 43 D. V. Yelnikov, A. N. Khanaychenko Material and method Biological analysis of 214 wild BST specimens caught during scientific ichthyological survey in Sev- astopol coastal area (North-Western shelf off Crimea — NWC, Black Sea) in 2008 during their natural spawning period (March–June) at the depths 40–70 m was carried out. Among the sampled fish, 12 mature specimens of both genders, of the standard length varying from 36.5 to 51.0 cm without any observed abnor- malities in skeleton structure were selected to describe standard normal morphological features of cephalic skeleton of adult BST. The cranium was separated from the soft tissues by processing the fish skulls in the hot water, followed by washing in running tap water. The bones were disarticulated after natural maceration in a tap water in closed vessels, followed by careful washing in running tap water. Otoliths for fish aging were extracted through the large occipital foramen prior to disarticulation of the bones. BST age was determined by counting the annual rings on the medial side of the sagittal otolith under the microscope MBS–10 under transmitted light. Selected for description of morphological features of BST fishes were aged after otolith analysis from 4 to 8 years old. Determination of specific bones and their spatial location in the skeletons of investigated specimens was carried out basing on the standard description of the bony fish skeleton after Gourtovoy et al. (1976). General- ized layout of standard BST cephalic skeleton was proposed after specialized scheme of the skeleton structure of Atlantic Pleuronectidae (Diaz de Astarloa, 2005). The nomenclature used to present the generalized scheme of the adult BST cephalic skeleton features followed the up-to-date descriptions of a multi-species anatomy ontology for teleost fishes (Whetzel et al., 2011) and after Dictionary of ichthyology (Coad, McAllister, 2007) and specialize descriptions of the flatfish cephalic skeleton (Chanet, 2003; Diaz de Astarloa, 2005). Results Cephalic skeleton (cranium, or skull) of BST as in all Teleosts consists of two main parts: Neu roc r an i um (axial skeleton of the head, or the braincase) and Vi s c e r o - c r an i um (syn. Sp l anchnoc r an i um, or facial skeleton of the head). Neu roc r a - Fig. 1. Structure of Neurocra- nium of the Black Sea turbot: À — dorsal view; B — left lat- eral view; C — right lateral view. 1 — parasphenoideum; 2 — vomer; 3 — basioccipitale; 4 — exoccipitale laterale; 4a — intercalare; 5 — supraoccipitale; 6 — parietale; 7 — frontale; 8 — ethmoideum laterale; 9 — me- sethmoideum; 10 — epioticum; 11 — pterosphenoideum; 12 — pteroticum; 13 — sphenoticum; 14 — prooticum. Original draw- ing of D. V. Yelnikov edited by A. V. Drapun. Ðèñ. 1. Còðîåíèå ÷åðåïíîé êîðîáêè ÷åðíîìîðñêîãî êàë- êàíà (P. ò. maeotica): À — âèä ñâåðõó, B — âèä ñëåâà, C — âèä ñïðàâà. 1 — parasphenoideum; 2 — vomer; 3 — basioccipitale; 4 — exoccipitale laterale; 4 a — intercalare; 5 — supraoccipitale; 6 — parietale; 7 — frontale; 8 — ethmoideum laterale; 9 — me- sethmoideum; 10 — epioticum; 11 — pterosphenoideum; 12 — pteroticum; 13 — sphenoticum; 14 — prooticum. Îðèãèíàëü- íûé ðèñóíîê Ä. Â. Åëüíèêî- âà. Õóäîæíèê-îôîðìèòåëü — À. Â. Äðàïóí. Unauthenticated Download Date \| 12/15/16 7:15 PM 44Morphological Features of Cephalic Skeleton of the Adult Black Sea Turbot… n ium is composed of the skull roof (or cranial dome) and the base of the braincase; sphenoid, occipital (syn. basicranial), orbital, olfactory (syn. ethmoid) and otic regions. V i s c e r o c r an i um is presented by visceral arches: the jaws, hyoid and gill arches. The general peculiarity of the adult BST is the left-right asymmetry of cephalic skeleton re- sulted from remodeling induced by metamorphosis. S t r u c t u r e o f BST Neu roc r an i um At the base of Neu roc r an i um (fig. 1: 1), two unpaired bones: parasphenoideum (fig. 1: 1) and vome r (fig. 1: 2) are located. Pa r a s pheno i d eum playing the role of the main beam of the cranium, stretches along the entire length of Neu roc r an i um , and its rostrum is fastened tightly to vome r under the eye-socket and to the main oc- cipital bone, b a s i o c c i op i t a l e in its back part (fig. 1: 3 — ba s i o c c i op i t a l e). On both sides of p a r a s pheno i d eum , its plate-like extensions are connected at their mid- part with f r on t a l e and p t e r o spheno i d eum, postero-laterally it articulates with p r oo t i c um and, posteriorly with b a s i o c c i p i t a l e. In its rostral part, p a r a s phe - no i d eum is bended slightly towards the ocular part of the body that is common to all flatfishes. In Psetta (or syn. Scophthalmus) genus, BST including, the right eye, as a rule (excluding abnormal, reversed specimens), migrates during metamorphosis to the left side of the body and, correspondingly, p a r a s pheno i d bends, with minor variations, towards the left side. Vome r, located in front of p a r a s pheno i d eum is significantly smaller than the latter and is extended in its rostral part. Lateral thickenings of v ome r are joined with the upper jaw, and on its lower part the teeth are located. Pa r a s pheno i d eum is the basis for several Neu roc r an i um regions — sphe- noid region, where conjoining with p t e r o spheno i d eum it forms the basis and the side walls of the middle part of the cranium; conjoining with e t hmo ida l e l a t e r a l e, me s e t hmo ideum and vome r it provides support to olfactory region; conjoining with f r on t a l e, it supports the orbital region. Occipital (syn. basicranial) region of the head is formed by b a s i o c c i op i - t a l e, e xoocc i p i t a l e and s up r aocc i p i t a l e (fig. 1: 3–5) delimiting the walls of the principal occipital foramen (f o r amen occ i p i t a l e magnum). As a result of asymmetric remodeling and body torsion of BST during metamorpho- sis, the main unpaired occipital bone, forming the floor of foramen magnum, b a s i o c - c i p i t a l e (fig. 1, B: 3 and fig. 1, C: 3) is shifted significantly from its longitudinal axis; it is attached to p a r a s pheno i d eum by its rostral extension, and its hind concave surface compatible with the surface of vertebrae centrum, serves a place of articulation with the first precaudal vertebra. Ventrally b a s i o c c i p i t a l e is linked to p a r a s phe - no i d forming p a r a s pheno i d -ba s i o c c i p i t a l e joint, anteriorly it articulates with e xoocc i p i t a l e. Above the dorsal surface of b a s i o c c i p i t a l e, and strongly articulated dorsally with it, two paired practically symmetrical, pyramidal-like from the lateral view, forming the lateral sides of foramen magnum, exoccipitals (e xoocc i p i t a l e, syn. = e xocc i p i - t a l e= occ i p i t a l e l a t e r a l e) — fig. 1, B: 4 and fig. 1, C: 4) are located. Together with b a s i o c c i p i t a l e, they form ba s i o c c i p i t a l e - e xocc i p i t a l e joint. At the place of this joint e xoocc i p i t a l e forms two overhanging processes. The right e xoocc i p i - t a l e could exceed the left one in size. Both have two large foramina allowing the pas- sage of two cranial nerves — glossofaryngeal and vagus. Anteriorly e xoocc i p i t a l e is bordered with p r oo t i c um, and with small membrane bone i n t e r c a l a r e laterally (fig. 1: 4a), the latter found in-between these larger bones. Dorsally e xoocc i p i t a l e borders the ventral surface of e p i o t i c um. Unpaired upper occipital bone (fig. 1: 5 — sup r aocc i p i t a l e), located at the dor- sal part of the occipital region of the cranium, postero-ventrally articulates with exooc- Unauthenticated Download Date \| 12/15/16 7:15 PM 45 D. V. Yelnikov, A. N. Khanaychenko cipitales, and completes the bony ring around the f o r amen magnum, forming its dorsal margin. On its top, s up r aocc i p i t a l e forms the high crest-like apophysis (fig. 1: 5), stretched forward far beyond the lower part of the bone thus separating the right and the left parietal and frontal bones. Front lateral edges of s up r aocc i p i t a l e are joined with the small, slightly asymmetrical paired parietal bones of otic region (p a r i e t a l e — fig. 1: 6) located dorsally in the back of the skull. The latter looks like trapezoid bony plate with thickened outer edges, and the smaller left stretches forward the larger right one. These bones cover the top of the head over otic region. Pa r i e t a l e is articulated to f r on t a l e anteriorly, and extensively overlaps the edge of p t e r o t i c um alongside the posterolateral edge of the Neu roc r an i um. Orbital region Axial torsion of BST Neu roc r an i um (as a rule to the left, up to 90o) takes place simultaneously with the migration of the right eye to the left side during metamorphosis, and leads to significant modification (relocation and deformation), and as a result, to asymmetry of all bones of the periorbital area: the frontal bones and the bones of ol- factory region. Frontal bones (f r on t a l e — fig. 1: 7), which form the orbital area, are altered most significantly during metamorphosis and are transformed into asymmetrical cup-like spongy bone plates with the crescent-shaped processes in their front lower part. The right f r on t a l e (fig. 1, B: 7) undergoes significant displacement and torsion along its longitudinal axis: it is deeper and more elongated lengthwise; its crescent-shaped process is longer, narrower and flatter than that of the left f r on t a l e, and sheathed in the latter. The crescent-shaped process of the right f r on t a l e together with the bones me s e t hmo ideum and the right e c t o e t hmo ideum forms the posterior border of the orbit of the right eye. The left f r on t a l e (fig. 1, C: 7) is wider and shorter than the right one. The lami- nated, large, crescent-shaped process located in the lower part of the left f r on t a l e is connected anteriorly with the left lateral olfactory bone (e c t o e t hmo ideum), and separates the left and the right eye-sockets. The left eye is not bordered by its own real eye-socket; it is located directly on the jaw muscles, outside the axial skull (n eu roc r a - n i um). The plexus of the bones of V i s c e r o c r an i um forms the bottom of the orbit of the left eye. Lacrimal bone (l a c r ima l e, fig. 2: 24) is a small bone lamina located at the front edge of the orbital foramen, above the junction of two visceral bones — pa l a t i num (fig. 2: 21) and e c t op t e r y g i o i d eum (fig. 2: 22). The group of small orbital bones (i n f r ao r b i t a l i a ) is typical for the orbital part of Neu roc r an i um in bony fishes (Teleosts) with lateral symmetry of the body (Gurtovoy et al., 1976). Most of these bones except one of them — small l a c r ima l e, are indistinguishable in the skeleton of adult BST. Other i n f r ao r b i t a l i a apparently fused with the neighbouring bones, either been reduced or lost in early ontogenesis. Together with the bones of olfactory region frontal bones cover the most of BST skull and frames (peri)orbital region of the skull. O l f a c t o r y , o r e t hmo id r e g i on is located directly above vome r and is at- tached to it. It is formed by paired lateral olfactory bones: e c t e t hmo ideum (= e t h - mo ideum l a t e r a l e = pa r e t hmo ideum = p l eu r e t hmo ideum = = p r a e - f r on t a l e = e x e t hmo ideum (Dias de Astarloa, 2005; McAlister, 2007) — fig. 1: 8) — left (fig. 1, B: 8) and right (fig. 1, C: 8) and unpaired me s e t hmo ideum (fig. 1: 9). Lateral ethmoids lie deeply in front of the orbits, and present the most signifi- cant size asymmetry among the bones of BST ethmoid region. They undergo significant transformation during metamorphosis and differ from each other both in shape and size. Contrary to BST frontal bones, the BST right e t hmo ideum (fig. 1, C: 8) is robust, Unauthenticated Download Date \| 12/15/16 7:15 PM 46Morphological Features of Cephalic Skeleton of the Adult Black Sea Turbot… considerably (almost twice) larger than the left one (fig. 1, B: 8), and has a massive wing-like expanded process that is connected with the right f r on t a l e. Unpaired intermediate olfactory bone (fig. 1: 9 — me s e t hmo ideum) is located in the front part of the skull (anterior part of the braincase) between two lateral olfac- tory bones. It is a bony bilobed plate curved in longitudinal direction. At the place of its junction with the lateral olfactory bones (fig. 1: 8 — e c t o e t hmo ideum) it forms the inner walls of olfactory capsules. Olfactory capsules are located at the top of the head, in front of f r on t a l e, between the bones of Sp l anchnoc r an i um and me s e t hmo i - d eum. From above they are covered with the paired nasal bones (na s a l i a, fig. 2: 20) separated by me s e t hmo ideum. Three bones (unpaired me s e t hmo ideum and two paired e c t o e t hmo ideum) are connected together in front of the skull and are joined with the upper part of v ome r. O t i c r e g i on (or auditory capsule) is located in postero-lateral of BST Neu ro - c r an i um and is comprised of series of paired bones located at each lateral side of the head, participating in formation of the walls of the otic capsule and supporting the vesti- bulo-auditory system. Most of these paired bones present only minor signs of asymmetry. Upper elements of otic capsule, paired bones (fig. 1: 10 — ep i o t i c um = = ep i - o c c i p i t a l e), locate at the back top of Neu roc r an i um and look like sharply outlined irregular frusto-conical tubercles. The ep i o t i c um is surrounded by the following bones: e xocc i p i t a l e (fig. 1: 4) (ventrally), s up r aocc i p i t a l e (fig. 1: 5) (laterally, along its axis), p a r i e t a l e (fig. 1: 6) (dorso-laterally), p t e r o t i c um (fig. 1: 12) (laterally). The shoulder girdle joins to ep i o t i c um through po s t t empo r a l e (not shown on the scheme of the BST cranium, since these bones are included into the shoulder girdle scheme). Together with p r oo t i c um and p t e r o t i c um it forms the auditory capsule encasing the semicircular canals of the inner ear where 3 pairs of otoliths are located. P t e r o spheno i d eum (fig. 1: 11) — is a paired, slightly asymmetric flat and hexagonal-shaped bone plate (the left — more circular and the right — more rectangu- lar) belonging to several regions: to sphenoid region — lying on the lateral expansion of p a r a s pheno i d and forming p a r a s pheno i d eum-p t e r o spheno i d eum joint as part of the lateral borders of the cranium; to orbital region joining with f r on t a l e ante- riorly; to otic region — sutured to p r oo t i c um posteriorly and, thus forming p roo t i - cum-p t e r o spheno i d eum joint and, dorso-posteriorly joins with s pheno t i cum. Foramina of p t e r o s pheno i d eum give the route to facial nerves. Large dense wing-shaped triangular paired bones (p t e r o t i c um fig. 1: 12) occupy the forepart of the otic region. Practically symmetrical, still the body of the left p t e r - o t i c um is insignificantly larger and has wider but shorter wing-like process than the right one. Together with p a r i e t a l e (fig. 1: 6) which locates immediately above p t e r - o t i c um and slightly overlaps it, they present very special finger- like curved surface which forms the postero-lateral border of the Neu roc r an i um . From the front the wing-like process of p t e r o t i c um is connected to f r on t a l e. P t e r o t i c um is sutured to postero-lateral surface of p r oo t i c um (fig. 1: 14) and, anteriorly to posterior surface of s pheno t i cum (fig. 1: 13), with e xocc i p i t a l e (fig. 1: 4) posteriorly, and postero- ventrally at a very restricted surface, with i n t e r c a l a r e (fig. 1: 4a), and is covered by ep i o t i c um (fig. 1: 10) dorso-posteriorly. Spheno t i cum syn. au t o spheno t i c (fig. 1: 13) — is a paired bone practically untouched by asymmetry resulted from the torsion of the cranium in BST. It has typi- cal saddle-like form with wing-like processes extended on both sides from its central part. Separating the orbital region from the otic region of the cranium, it is articulat- ed with p t e r o spheno i d eum antero-ventrally, with f r on t a l e dorso-medially, with p t e r o t i c um dorso-posteriorly and with p r oo t i c um ventrally; dorso-posterior part of s pheno t i cum very shortly sutures with p a r i e t a l e. Together with the latter, s phe - no t i cum participate in formation of the articulation fossa of h yomand i bu l a r e. Unauthenticated Download Date \| 12/15/16 7:15 PM 47 D. V. Yelnikov, A. N. Khanaychenko Considerable part of otic department is presented by the large paired p roo t i c um (fig. 1: 14), the bone shaped as improper square, or squared flower with a non-ossified cartilage area in its central part and perforated by foramen for trigeminal nerve. Thin foliated delicate posterior surface of p r oo t i c um is sutured ventrally with the thin ante- rior surfaces of b a s i o c c i p i t a l e forming together with it the chamber where the sagit- tal otoliths are located. Postero-dorsally p r oo t i c um is sutured densely with p t e r o t i - cum . Antero-dorsally p r oo t i c um is connected to the ventral part of s pheno t i cum, and at the boundary of their joining the articulation of h yomand i bu l a r e is located. Antero-ventrally p r oo t i c um is articulated with the lateral extensions of p a r a s phe - no i d, and anteriorly it borders with p t e r o spheno i d eum; while dorso-posteriorly it borders with e xooc i p i t a l e and, slightly with i n t e r c a l a r e (fig. 1: 4à). The smallest bone of the otic region, cone-form, i n t e r c a l a r e (syn. op i s t ho t i - cum = pa rocc i p i t a l e (fig 1: 4a) is bordered with anterior surface of e xoocc i p i - t a l e and posterior surface of p t e r o t i c um, and, shortly, with p r oo t i c um, serving the posterior wall of otic capsule. I n t e r c a l a r e is often not easily found and often lost among the Neu roc r an i um bones of BST in case of absence of preliminary macera- tion of the skull prior to disarticulation of the bones. Small, slightly asymmetrical paired parietal bones of otic region (p a r i e t a l e — fig. 1: 6) locate dorsally in the back of the skull. Those are trapezoid bony plates with thick- ened outer edges: the lighter and smaller left is stretched slightly forward than the larger right one; and both articulating medially cover the top of the head over otic region, posteriorly to s up r aocc i p i t a l e. Pa r i e t a l e is articulated with f r on t a l e from ahead and extensively overlaps the edge of p t e r o t i c um alongside the posterolateral edge of the Neu roc r an i um. Structure of BST Splanchnocranium (branchiocranium, or visceral cranium) All bones of Sp l anchnoc r an i um including the jaws, hyoid and branchial re- gions (fig. 2–4), are paired in Teleosts including Scophthalmidae. What is distinctive for the visceral skeleton of BST, as for Psetta (Scophthalmus) genus in general, is that the bones of the blind side (normally, the right one) have lesser (minor) spatial curvature than the bones of the ocular side (normally, the left one). In other aspects (the differ- ences in location and size characteristics of the left-right bones) are almost negligible. Mand i bu l a r (j aw s) a r ch (o r a r cu s mand i bu l a r i s ) is represented by the upper and lower jaws (upper and lower jaw elements). Lowe r j aw . Dentary bone (d en t a l e, fig. 2: 17) lies in the rostral part of the lower jaw and bears numerous small teeth. By its lower and more elongated process, d en t a l e attaches to a r t i c u l a r e. A r t i c u l a r e (fig. 2: 18) is located in the posterior part of the mandible. Upper jaw bone — quad r a t um (fig. 2: 23) is adjustably fixed in articular bursa of the rear top of a r t i c u l a r e. Small an gu l a r e (fig. 2: 19) terminates the lower jaw. It almost adheres to the inner surface of the lower rear edge of a r t i c u - l a r e. The asymmetry of the bones of the lower jaw arch is negligible; the bones of the right (blind) side — den t a l e, a r t i c u l a r e and angu l a r e — are slightly more massive than similar bones of the left (ocular) side. Uppe r j aw is articulated with the lower jaw through the primary bones of the upper jaw, quadrates (quad r a t um, fig. 2: 23). Despite the name, those are triangle shaped osseous trabecula with a comb-like crest at their ventral side. The articular sur- face of quad r a t um epiphysis is located at the apex of the triangle of quad r a t um; the bony lamina of quad r a t um is stretched between two robust beams of almost isoscales triangle. The left quad r a t um is slightly wider and shorter than the right one. The up- per bound of quad r a t um is attached to the outside of e c t op t e r y go i d eum (fig. 2: 22). Ectopterygoids are the narrow curved paired boomerang-like bones (the right one Unauthenticated Download Date \| 12/15/16 7:15 PM 48Morphological Features of Cephalic Skeleton of the Adult Black Sea Turbot… slightly stronger than the left one) connected by its interior surface to me t ap t e r y go i - d eum; from their front side they are connected with p a l a t i num. Hook-like bones — pa l a t i num (fig. 2: 21), the right slightly larger than the left one, are movably attached by their anterior part to Neu roc r an i um through vome r articulation. The interweav- ing of three pterygoid bones forms the left eye-socket. The thin round lamina of internal pterygoid bone — en top t e r y go i d eum (= mesop t e r y go i d eum, fig. 2: 25) is fixed to the hind part of e c t op t e r y go i d eum. Posterior pterygoid bone (me t ap t e r y - go i d eum, fig. 2: 26) is located between en t op t e r y go i d eum and quad r a t um. Secondary upper jaw is formed by two pairs of bones: premaxillary (fig. 2: 15 — p r a emax i l l a r e = i n t e rmax i l l a r y = s u rmax i l l a r y = b imax i l l a r y) and maxil- lary (fig. 2: 16 — max i l l a r e). Premaxillary bones located at the tip of upper jaw, are the thin curved laminas with several processes (ascending, articular, postmaxillary and caudal); both premaxillary bare small teeth. Maxillaries — are the curved laminas, ex- panded in their posterior part, with articular formations in their anterior part. Both pairs of bones of the secondary upper jaw are joined movably together by the rostral cartilage in one point of Neu roc r an i um above vome r . Insignificant asymmetry of the bones of the upper jaw of BST is the following: the bones of the ocular side of the body, espe- cially, max i l l a r e, are more robust and more curved than the bones of the blind side. Hyo i d r e g i on of Sp l anchnoc r an i um, lying between the jaws and gil l arches is represented by the upper branch, called otherwise dorsal hyoid branch, and the lower branch, otherwise known as ventral hyoid branch. The basis of the upper branch is h yomand i bu l a r e (fig. 2: 27) which serves as articulation of the jaws apparatus (suspension) to the otic department of n eu roc r a - n i um, connecting the lower jaw with the skull. The anterior part of h yomand i bu l a r e Fig. 2. Structure of mandibular arch of the Black Sea turbot (P. m. maeotica): 15 — praemaxillare; 16 — maxil- lare; 17 — dentale; 18 — articulare; 19 — angulare; 20 — nasale; 21 — palatinum; 22 — ectopteryogoideum; 23 — quadratum; 24 — lacrimale; 25 — entopteryogoideum; 26 — metapteryogoideum; 27 — hyomandibulare; 28 — symplecticum. Gill cover structure: 29 — praeoperculum; 30 — operculum; 31 — suboperculum; 32 — interoperculum. Original drawing of D. V. Yelnikov edited by A. V. Drapun. Ðèñ. 2. Ñòðîåíèå ÷åëþñòíîé äóãè ÷åðíîìîðñêîãî êàëêàíà (P. m. maeotica): 15 — praemaxillare; 16 — max- illare; 17 — dentale; 18 — articulare; 19 — angulare; 20 — nasale; 21 — palatinum; 22 — ectopteryogoideum; 23 — quadratum; 24 — lacrimale; 25 — entopteryogoideum; 26 — metapteryogoideum; 27 — hyomandibulare; 28 — symplecticum. Ñòðîåíèå æàáåðíîé êðûøêè: 29 — praeoperculum; 30 — operculum; 31 — suboperculum; 32 — interoperculum. Îðèãèíàëüíûé ðèñóíîê Ä. Â. Åëüíèêîâà. Õóäîæíèê-îôîðìèòåëü — À. Â. Äðàïóí. Unauthenticated Download Date \| 12/15/16 7:15 PM 49 D. V. Yelnikov, A. N. Khanaychenko connects to pterygoids and its posterior part is articulated with the bones of operculum. Paired bone s ymp l e c t i c um (fig. 2: 28) is attached to the lower part of h yoman- d i bu l a r e. It is a strong thin elongated arcuately curved, bilaterally sharpened bone expanded in the curved part. The lower part of s ymp l e c t i c um binds tightly to the bones of the upper jaw, to the lower compacted edge of quad r a t um . Ventral hyoid arch (fig. 3) is attached to hyomand i bu l a r e through i n t e r hy a l e (fig. 3: 33), a relatively small, elongated bone thickened ventrally, attached by cartilage to the lower bone — ep i hy a l e (fig. 3: 34). This triangular-like bone at the upper end of hyoid arch together with c e r a t ohya l e (fig. 3: 35) forms a tight entity with a barely noticeable crest-like joint in its middle part. Wide connecting suture (in the form of a crest-lock) between these two bones widen towards the lateral parts and generates the triangular cartilage densification at its edges (fig. 3: 34, 35). The paired hypohya l e (fig. 3: 36) composed of upper and lower tightly fused parts, is joined with the long process of the anterior part of c e r a t ohya l e. Below, under the branchial apparatus, the unpaired bone u rohya l e (fig. 3: 37, syn. b a s i b r anch i o s t e g a l e) is fastened to a group of major branchial bones and the bones of shoulder girdle with the help of the tendon. Another unpaired bone of elongated form with a cone-shaped cartilage at its an- terior edge (b a s i h y a l e , syn. g l o s s ohya l e, fig. 3: 38) is located at the front end of the distal part of the hyoid arch. Hyoid arches are movably attached to the recesses in posterior part of b a s i h y a l e. Branchiostegal rays, or r ad i i b r anch i o s t e g i i (fig. 3: 39) branch off e p i h y a l e and c e r a t ohya l e and serve the basis for branchial membrane. All of the above men- tioned bones of hyoid arch (with exception of b a s i h y a l e ) at both sides of the body are paired and practically symmetrical. Minor differences in all paired bones of hyoid arch on the right and left sides of the body are found in various density in some parts of certain bones and in directions of their bending and jointing. G i l l a r ch s uppo r t s ( b r anch i a l a ppa r a t u s ) The central part of BST gill apparatus is comprised of three main/ principal axial unpaired branchial bones (b a s i b r anch i a l e = copu l a, fig. 4: Bb.1–Bb.3), tightly held together by means of strong cartilaginous joints. Paired symmetrical gi l l arches are attached to the principal branchial bones. Three pairs of the upper branchial arches are attached, respectively, to each side of three b a s i b r anch i a l e. The short bones (hy - Fig. 3. Structure of hyoid arch of the Black Sea turbot (P. m. maeotica): 33 — interhyale; 34 — epihyale; 35 — ceratohyale; 36 — hypohyale; 37 — urohyale; 38 — basihyale; 39 — radii branchiostegii. Original drawing of D. V. Yelnikov edited by A. V. Drapun. Ðèñ. 3. Ñòðîåíèå ãèîèäíîé äóãè ÷åðíîìîðñêîãî êàëêàíà (P. m. maeotica): 33 — interhyale; 34 — epihyale; 35 — ceratohyale; 36 — hypohyale; 37 — urohyale; 38 — basihyale; 39 — radii branchiostegii. Îðèãèíàëüíûé ðèñóíîê Ä. Â. Åëüíèêîâà. Õóäîæíèê-îôîðìèòåëü — À. Â. Äðàïóí. Unauthenticated Download Date \| 12/15/16 7:15 PM 50Morphological Features of Cephalic Skeleton of the Adult Black Sea Turbot… pob r anch i a l e, f i g . 4: Hb.1–Hb.3), are adjustably fastened to the first three branchial arches. The fourth and fifth gill arches are attached through cartilage to the ventral inferior end of the last basi- branchiale. Each branchial arch includes c e r a - t ob r anch i a l e (fig. 4: Cb.1–Cb.5) — light elongated bones with the grooves on their distal part which contain blood vessels coming from the b r anch i o s t e - g a l r a y s. The fifth c e r a t ob r anch i - a l e bears the small cloves. Paired up- per gill bones (ep i b r anch i a l e, fig. 4: Eb.1–Eb.4), each pair having its proper distinctive form, are located on the first four branchial arches posteriorly c e r a - t ob r anch i a l e. Pharyngobranchials (pha r yngob r anch i a l e fig. 4: Fb.1– Fb.4) are located in the distal part of the first four branchial arches. Fused togeth- er, they form the pharyngeal apparatus. All pharyngobranchials, except for the first pair, are smooth and elongated, and have the bony shields bearing the teeth. Gil l arches are covered by the gi l l cover, thin solid plate consisted of four dermal paired bones of opercu- lum series (fig. 2: 29–32). The surface of the paired sickle-shaped preopercular bone (p r a eope r cu l um, fig. 2: 29) is tightly attached to posterior external part of the hyomandibular bone. The upper edge of the gill cover is formed by the upper side of the anterior edge of opercular bone (ope r cu l um, fig. 2: 30) movably attached in the joint of hyomandibular bone. From below ope r cu l um adjusts to a small flat bone s ubope r cu l um (fig. 2: 31). I n t e r o - pe r cu l um, (fig. 2: 32) joins along its perimeter with the other three gill cover bones and is connected with angu l a r e of the lower jaw by means of the tendons. P r a eo - pe r cu l um is partly overlying i n t e r ope r cu l um (by its horizontal part) and (by its vertical part). Left-right asymmetry of the gill apparatus is weakly expressed — mainly in insignificant difference in sizes and shapes of paired bones. Conclusions The most typical morphological features of the cranial skeleton of the Black Sea kalkan, as in all species of Pleuronectiformes is the asymmetrical structure of the left and right sides of n eu roc r an i um forming during metamorphosis. Under normal con- ditions, during the metamorphosis, the right eye totally migrates to the left side, and synchronously with the right eye migration, the bones restructure, and asymmetry in practically all the paired bones of neurocranium develops. Among all the bones of the BST Neurocranium, the bones of periorbital region acquired most significant asymmetry as a result of early eye migration and cranial skeleton torsion towards the ocular left side Fig. 4. Gill arch structure of the Black sea turbot (dorsal view, left side). Bb.1–Bb.3 — basibranchiale; Cb.1– Cb.5 — ceratobranchiale; Eb.1–Eb.4 — epibranchiale; Fb.1–Fb.4 — pharyngobranchiale; Hb.1–Hb.3 — hypobranchiale. Original drawing of D. V. Yelnikov edited by A. V. Drapun. Ðèñ. 4. Ñòðîåíèå æàáåðíîé äóãè ÷åðíîìîðñêîãî êàëêàíà (âèä ñâåðõó, ëåâàÿ ñòîðîíà òåëà). Bb.1– Bb.3 — basibranchiale; Cb.1–Cb.5 — ceratobranchiale; Eb.1–Eb.4 — epibranchiale; Fb.1–Fb.4 — pharyn- gobranchiale; Hb.1–Hb.3 — hypobranchiale. Îðè- ãèíàëüíûé ðèñóíîê Ä. Â. Åëü íèêîâà. Õóäîæíèê- îôîðìèòåëü — À. Â. Äðàïóí. Unauthenticated Download Date \| 12/15/16 7:15 PM 51 D. V. Yelnikov, A. N. Khanaychenko of the body and main deformations took place in anterior part. The most considerable differences are observed for the paired bones f r on t a l e and e c t o e t hmo ideum. Sev- eral bones of cephalic skeleton — group of orbital bones i n f r ao r b i t a l i a (except l a c - r ima l e), as well as bones common to most of bony fishes, such as o r b i t o s pheno i - d eum, and ba s i s pheno i d eum, were not found in BST, probably being reduced, or completely fused with the other bones, or lost as in Gadidae. Left-right asymmetry in the Sp l anchnoc r an i um in BST is less expressed than asymmetry of the Neurocranium. The asymmetry of the right and left bones of the mandibular apparatus appears in minor dimensional and density characteristics. Bones of hyoid arch are practically symmetrical. Authors are grateful to designer Anna Drapun for significant help in edition of original drawings; Dr. Yu- nia Bityukova, Dr. Michael Chesalin and Dr. Bruno Chanet for scientific consultancies; Dr. Vitaly Giragosov for the help in fish ageing and scientific consultancies. The manuscript preparation was partly supported by FP7 PERSEUS project (GA 287600) and Project 0113U003601 of National Ukrainian Academy of Sciences. References Chanet B. Interrelationships of scophthalmid fishes (Pleuronectiformes: Scophthalmidae). Cybium. — 2003. — 27, N 4. — Ð. 275–286. Coad B. W., McAllister D. E. Dictionary of Ichthyology. — 2007. — http://www.briancoad.com/Dictionary/ Complete%20Dictionary.htm. Diaz de Astarloa J. M. Osteologia craneal comparada de tres especies de lenguado del genero Paralichthys (Pleuronectiformes, Paralichthyidae) del Atlantico suroccidental // Rev. chil. hist. nat. — 2005. — 78, N 3. — P. 343–391. Gourtovoy N. N., Matvejev B. S., Dzerzhinsky F. Y. Practical zootomy of the vertebrates (lower vertebrates, agnathous, pisces). Uchebnoye posobiye dlya biologicheskikh spetsial’nostey universtitetov. — Moskva : Vysshaya shkola, 1976. — P. 302–324. — Russian : Ãóðòîâîé Í. Í., Ìàòâååâ Á. Ñ., Äçåðæèíñêèé Ô. ß. Ïðàêòè÷åñêàÿ çîîòîìèÿ ïîçâîíî÷íûõ (íèçøèå õîðäîâûå, áåñ÷åëþñòíûå, ðûáû). Kalinina E. M. On anatomy of the Black Sea flatfish (Pleuronectiformes) // Trudy Sevastopolskoy biolog- icheskoy stantsii. — 1959. — 12. — P. 319–327. — Russian : Êàëèíèíà Ý. Ì. Ê àíàòîìèè ÷åðíîìîðñêèõ êàìáàëîîáðàçíûõ (Pleuronectiformes). Khanaychenko A. N., Giragosov V. E., Yelnikov D. V., Danilyuk O. N. Pigmentation anomalies in the Black Sea turbot Psetta (= Scophthalmus) maxima maeotica (Pleuronectiformes: Scophthalmidae) // Morskiy Ekologichny Zhurnal. — 2008. — 7, N 2. — P. 87–95 — Russian : Õàíàé÷åíêî À. Í., Ãèðàãîñîâ Â. Å., Åëüíèêîâ Ä. Â., Äàíèëþê Î. Í. Àíîìàëèè ïèãìåíòàöèè ÷åðíîìîðñêîé êàìáàëû êàëêàíà Psetta (= Scophthalmus) maxima maeotica (Pleuronectiformes: Scophthalmidae). Whetzel P. L., Noy N. F., Shah N. H. et al. BioPortal: enhanced functionality via new Web services from the National Center for Biomedical Ontology to access and use ontologies in software applications // Nucleic Acids Res. — 2011. — doi: 10.1093/nar/gkr469. Received 3 August 2012 Accepted 1 October 2013 Unauthenticated Download Date \| 12/15/16 7:15 PM

Morphological Features of Cephalic Skeleton of the Adult Black Sea Turbot (Kalkan) Scophthalmus maximus var. maeotica (Pleuronectiformes, Scophthalmidae)

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