Diversity of dermal denticle structure in sharks: Skin surface roughness and three‐dimensional morphology

Shark skin is covered with numerous placoid scales or dermal denticles. While previous research has used scanning electron microscopy and histology to demonstrate that denticles vary both around the body of a shark and among species, no previous study has quantified three‐dimensional (3D) denticle structure and surface roughness to provide a quantitative analysis of skin surface texture. We quantified differences in denticle shape and size on the skin of three individual smooth dogfish sharks (Mustelus canis) using micro‐CT scanning, gel‐based surface profilometry, and histology. On each smooth dogfish, we imaged between 8 and 20 distinct areas on the body and fins, and obtained further comparative skin surface data from leopard, Atlantic sharpnose, shortfin mako, spiny dogfish, gulper, angel, and white sharks. We generated 3D images of individual denticles and measured denticle volume, surface area, and crown angle from the micro‐CT scans. Surface profilometry was used to quantify metrology variables such as roughness, skew, kurtosis, and the height and spacing of surface features. These measurements confirmed that denticles on different body areas of smooth dogfish varied widely in size, shape, and spacing. Denticles near the snout are smooth, paver‐like, and large relative to denticles on the body. Body denticles on smooth dogfish generally have between one and three distinct ridges, a diamond‐like surface shape, and a dorsoventral gradient in spacing and roughness. Ridges were spaced on average 56 µm apart, and had a mean height of 6.5 µm, comparable to denticles from shortfin mako sharks, and with narrower spacing and lower heights than other species measured. We observed considerable variation in denticle structure among regions on the pectoral, dorsal, and caudal fins, including a leading‐to‐trailing edge gradient in roughness for each region. Surface roughness in smooth dogfish varied around the body from 3 to 42 microns.     

<Emphasis Type="Italic">Icriodus marieae</Emphasis>, a new icriodontid conodont species from the Middle Devonian

A new conodont species, Icriodus marieae, is described from pelagic limestone beds of the Carnic Alps (Austria). Specimens are obtained from the upper part of the Valentin Formation (Central Carnic Alps) and range from the latest Eifelian to middle Givetian. Significantly differing from other icriodontid conodonts is that the icriodontan element of the new species develops only three denticles on either lateral denticle row, which are constricted to the central part of the element. The anterior part of the element is free of lateral row denticles and consists of two to four denticles, which have a fan-shaped outline in lateral view. The anterior part as well as the posterior part (consisting of cusp and two to three pre-cusp denticles) is higher than the denticles of the central part of the element. Shape analysis confirms that the parameters chosen for landmarks (element size relation and denticle setting) show little variation between different specimens.     

Patent infections of Ascaris suum in pigs: effect of previous exposure to multiple, high doses of eggs and various treatment regimes.

Fifty-four crossbred, 4-week-old pigs divided into nine equal groups were used to test whether multiple inoculations with high numbers of A. suum eggs with or without anthelmintic would result in patent infections. All pigs exposed to multiple prechallenge inoculations of 500, 1000, 2000, 5000, 10,000 and 20,000 and challenged orally 2 weeks later with 10,000 eggs harboured adult worms. When prechallenge infections were removed by pyrantel tartrate treatment the animals were more susceptible to challenge than controls not previously exposed to infections. The same drug used from 2 days before until 10 days after the last prechallenge infection eliminated that effect. Pigs subjected to the same multiple egg dosing regimen but given feed containing fenbendazole immediately before, during and for 10 days after multiple dosing developed significantly more adult intestinal worms after challenge than any other group. These worms were, however, significantly shorter than those that developed in any group of pigs. Adult worms from all these groups produced eggs that after embryonation were infective to mice.     

Morphology of the oral disc of Bufo bufo (Salientia: Bufonidae) tadpoles

The fully developed oral disc of the tadpole of Bufo bufo consists of dorsal and ventral labia bearing, respectively, two and three ridges bearing numerous horny denticles, a horny beak provided with jaw sheath serrations, and large lateral papillae that are borne by two cutaneous plicae. As development progresses toward metamorphosis, these structures gradually regress until they disappear. Each cusped clavate labial denticle adheres, by means of a thin peduncle, to a similar labial denticle fixed in the lip and formed by a group of three or four cells that keratinize gradually and thus present remarkable differences in their morphology. Once all the cells of a group have been converted into horny tissue, the denticle sheds and is replaced by the underlying one. The beak serrations also are horny structures; each consists of a columnar band of cells which undergoes a gradual keratinization. The horny cells that detach themselves at intervals, being replaced by those of the underlying anlagen. The labial denticles and the beak serrations keratinize in two distinct ways. In the former, the desmosomal filaments appear to play an important role whereas, in the latter, the keratin seems to be synthesized “ex novo” by the ribosomes.     

Anomalies in the expression of abdominal denticle belts of partial larvae produced by fragmentation of theDrosophila melanogaster egg

Summary Abnormal denticle belt patterns can occasionally be observed in abdominal belts of partial larvae obtained from egg fragments. The abdominal belts have the following features in common: 1) The number of denticles of an abdominal denticle belt may increase, depending on the space occupied by a distinct segment or the whole body region. The arrangement of the denticles in such enlarged belts is less regular than in normal belts. 2) Enlarged denticle belts are also found in the terminal segment of a fragment, or in the segment next to it when the larval pattern is interrupted by fragmentation. The denticle belt in the adjacent segment(s) may then be supressed. 3) All denticles in a belt (or part of a belt) are orientated posteriorly if the distance to the posteriorly adjacent belt (or part of a belt) is larger than normal, or if this denticle belt is suppressed. Conditions anterior to a segment do not seem to exert any influence on denticle orientation.     

The denticle surface of thresher shark tails: Three-dimensional structure and comparison to other pelagic species

Shark skin denticles (scales) are diverse in morphology both among species and across the body of single individuals, although the function of this diversity is poorly understood. The extremely elongate and highly flexible tail of thresher sharks provides an opportunity to characterize gradients in denticle surface characteristics along the length of the tail and assess correlations between denticle morphology and tail kinematics. We measured denticle morphology on the caudal fin of three mature and two embryo common thresher sharks (Alopias vulpinus), and we compared thresher tail denticles to those of eleven other shark species. Using surface profilometry, we quantified 3D-denticle patterning and texture along the tail of threshers (27 regions in adults, and 16 regions in embryos). We report that tails of thresher embryos have a membrane that covers the denticles and reduces surface roughness. In mature thresher tails, surfaces have an average roughness of 5.6 μm which is smoother than some other pelagic shark species, but similar in roughness to blacktip, porbeagle, and bonnethead shark tails. There is no gradient down the tail in roughness for the middle or trailing edge regions and hence no correlation with kinematic amplitude or inferred magnitude of flow separation along the tail during locomotion. Along the length of the tail there is a leading-to-trailing-edge gradient with larger leading edge denticles that lack ridges (average roughness = 9.6 μm), and smaller trailing edge denticles with 5 ridges (average roughness = 5.7 μm). Thresher shark tails have many missing denticles visible as gaps in the surface, and we present evidence that these denticles are being replaced by new denticles that emerge from the skin below.     

The characterization, replication and testing of dermal denticles of Scyliorhinus canicula for physical mechanisms of biofouling prevention

There is a current need to develop novel non-toxic antifouling materials. The mechanisms utilized by marine organisms to prevent fouling of external surfaces are of interest in this regard. Biomimicry of these mechanisms and the ability to transfer the antifouling characteristics of these surfaces to artificial surfaces are a highly attractive prospect to those developing antifouling technologies. In order to achieve this, the mechanisms responsible for any antifouling ability must be elucidated from the study of the natural organism and the critical surface parameters responsible for fouling reduction. Dermal denticles of members of the shark family have been speculated to possess some natural, as yet unidentified antifouling mechanism related to the physical presence of denticles. In this study, the dermal denticles of one particular member of the slow-swimming sharks, Scyliorhinus canicula were characterized and it was found that a significant natural variation in denticle dimensions exists in this species. The degree of denticle surface contamination was quantified on denticles at various locations and it was determined that the degree of contamination of the dorsal surface of denticles varies with the position on the shark body. In addition, we successfully produced synthetic sharkskin samples using the real skin as a template. Testing of the produced synthetic skin in field conditions resulted in significant differences in material attachment on surfaces exhibiting denticles of different dimensions.     

Acquired resistance to migrating larvae of Ascaris suum in young pigs by repeated drug-abbreviated infections

Cross-bred 3- and 8-wk-old pigs were used to test whether drug-abbreviated infections with Ascaris suum can stimulate acquired resistance to challenge. During the immunization period, both age groups of animals were infected with increasing numbers of A. suum eggs (500, 1,000, 2,000, 5,000, 10,000, and 20,000) at 7-day intervals while the pigs were receiving pyrantel tartrate in the feed. Two days after the last infective dose, animals were placed on unmedicated feed for 8 days and then challenged with 10,000 eggs. All pigs were killed 7 days after challenge, and milk spots on the livers and larvae recovered from the lungs were counted. Larval recoveries from lungs of the immunized animals were significantly smaller than those from the unimmunized animals in both age groups, suggesting that the pigs were capable of acquiring strong resistance to parasitic infections. In immunized animals, challenge infection did not contribute significantly to milk spot formation. The number of milk spots was significantly greater in the older animals, indicating that milk spot formation may be age related.     

Gill raker structure and development in indo-pacific anchovies (Teleostei: Engrauloidea), with a discussion of the structural evolution of engrauloid gill rakers

The postlarval development of gill raker denticles is described for the engrauloid (anchovy) genera Coilia, Lycothrissa, Setipinna, Thryssa, Stolephorus, and Encrasicholina based on scanning electron microscopy. The raker structure of adult Papuengraulis is also described. In the coiliid genera Coilia, Lycothrissa, Setipinna, Thryssa, and Papuengraulis, denticle development is not confined to particular region(s) of the raker. With few exceptions, the proliferation of denticles with growth is greatest along the upper raker edge; denticles are smaller and less dense on the raker faces and along the lower raker edge. Some Thryssa and Setipinna have a derived condition of denticle clustering along the upper raker edge. In Stolephorus and Encrasicholina, denticle development is confined to the upper raker half and includes the development of a single row of denticles along each raker face. A phylogenetic analysis of engrauloid raker structure, incorporating data from Bornbusch ( 88: Copeia 1988:174–182) and based on outgroup comparisons, indicates that for the Engrauloidea: (1) the pattern of denticle development shared by coiliids is plesiomorphic; and (2) the pattern of denticle development shared by Stolephorus, Encrasicholina, and most other engraulids is synapomorphic for the Engraulidae. There is no evidence that the studied coiliids Stolephorus and Encrasicholina are suspension feeders. The engraulid pattern of raker denticle development which is retained in suspension feeding engraulids of the genus Engraulis was thus derived before the derivation of suspension feeding in Engraulis. Comparative morphological and phylogenetic studies of clupeomorph raker structures and feeding behaviors can infer the historical origins of morphology-behavior associations, help define possible directions for analyses of raker denticle function, and thereby help elucidate the significance of structure-function couplings in the evolution of such clupeomorph trophic behaviors as suspension feeding. © 1992 Wiley-Liss, Inc.     

Relationship of hemolymph phenol oxidase and mosquito age in Aedes aegypti.

Monophenol oxidase (MPO) and diphenol oxidase (DPO) activity in hemocytes and cell-free plasma perfused from 7-, 14-, 21-, and 28-day-old Aedes aegypti mosquitoes were compared. A progressive decrease of enzyme activity was detected as mosquito age increased, and this decrease was significant in both hemocytes and cell-free plasma when mosquitoes were 28 days old as compared with that found in 7-day-old mosquitoes. There was no significant difference in total hemolymph protein as mosquito age increased. Although this decreased MPO and DPO activity might be partially responsible for the reduced immune response against filarial worms previously reported for older mosquitoes, other factors undoubtedly play a significant role.     

云南曲靖张家营一肺鱼齿板

<正> 本文记述的肺鱼齿板是1979年在云南进行野外工作时采获的。标本产自云南曲靖张家营东山中泥盆统曲靖组。登记号V6257 经观察这一标本很可能属于双翼鱼科(Dipteridae),代表一新属、新种。特征一保存不完整的齿板,冠面呈扇形。具9条齿脊,彼此近于平行,脊上具有数目不等的齿突,表面具有琺琅质层。齿谷表面粗糙并缺失琺琅质层。靠近齿板外缘内侧,在齿板冠面上有一浅槽。描述一件保存不完整的左下齿板,仅前侧具脊的部分被保存下来,而后中光滑的台面部分则缺失。齿板中等大小,呈扇形。保存部分的最大长度21毫米,最大宽度16毫     

Evaluation of pyrantel-tartrate abbreviated Ascaris suum infections for the development of resistance in young pigs against migrating larvae

Crossbred young pigs were used to test whether abbreviated infections with eggs of Ascaris suum can stimulate the acquisition of resistance to challenge. Weanling pigs from an Ascaris-free colony were kept free of A. suum until they were divided into groups at the age of 7-8 weeks. The experimental animals received pyrantel tartrate during the period when they were being exposed to increasing numbers of infective A. suum eggs and challenged 10 days after the last infective dose. Liver milk-spot counts and larval recoveries from the lungs indicated that the strongest resistance was acquired by the animals that received the drug continuously for 6 weeks while being exposed to six weekly infective egg doses. The data do not suggest any drug-related suppression of the resistance response to A. suum infection.     

Ascaris suum: development of intestinal immunity to infective second-stage larvae in swine

The development of protective immunity to Ascaris suum was examined in pigs naturally exposed to eggs on a contaminated dirt lot. Pigs became almost totally immune to second-stage larvae migrating from the intestines because few larvae from a challenge inoculum could be found in the lungs, and liver white-spot lesions (an immunopathologic response to migrating larvae) were absent. Blood from these pigs contained lymphocytes that responded blastogenically to larval antigens in vitro, while the serum contained antibody to larval antigens. Immunity was related to parasite exposure and not to the age of the host, and was not affected by the removal of adult A. suum from the intestines. Naturally exposed pigs responded to a variety of A. suum antigens with an immediate-type skin reactivity, and their intestinal mucosa contained relatively large numbers of mast cells and eosinophils. Other pigs were maintained on a dirt lot not contaminated with A. suum eggs and the effects of common environmental conditions on development of resistance to A. suum were studied. Resistance also developed in these pigs because 72% fewer larvae were detected in their lungs following a challenge exposure than in control pigs confined indoors on concrete floors and challenged similarly. This response was not expressed at the intestinal level, however, because their livers had numerous, intense white-spot lesions. To verify that the intestinal immunity that developed in pigs after natural exposure to A. suum was a direct result of homologous infection and not related to other stimuli encountered on a dirt lot, pigs maintained indoors on concrete floors, free from inadvertent helminthic infection, were inoculated orally with A. suum eggs daily for 16 weeks. Intestinal immunity was induced because larvae from a challenge inoculum were not detected in the lungs, and few white-spot lesions appeared on the livers of these pigs. Apparently, continual exposure of the intestinal mucosa to larvae eventually elicits the appropriate effector components necessary to prevent larval migration from the intestines.     

Effect of noradrenaline on oxygen consumption and tissue blood flow in young pigs

Metabolic rate and tissue blood flow were measured by indirect calorimetry and the radioactive microsphere technique, respectively, in anaesthetized pigs aged 10, 17, and 28 days during infusions of saline or noradrenaline. Noradrenaline produced a rise in oxygen consumption in all pigs but the magnitude of the response declined with age in Large White pigs (percent increase: 10 days old, 29.5 +/- 3.3; 17 days old, 15.3 +/- 3.2; 28 days old, 5.4 +/- 0.17) and was less in 10-day-old Minipigs (9.1 +/- 5.1). Regional blood flow per gram of tissue was similar for pigs of all ages and noradrenaline produced increases in flow to heart and skin, and small reductions in blood flow to skeletal muscle. Noradrenaline increased average blood flow to adipose tissue (four sites) by 21-fold in 10-day-old Large White pigs, but had less effect in 10-day-old Minipigs (9.5-fold) and in older Large White pigs (9-fold), where the effect on metabolic rate was diminished. It is calculated that adipose tissue could be responsible for the increase in metabolic rate during infusion of noradrenaline. Macroscopically there are no gross differences between various adipose tissue depots in the pig but those which showed the greatest response to noradrenaline correspond to areas where brown adipocytes have previously been identified by electron microscopy.     

Biological characterization of the skin of shortfin mako shark Isurus oxyrinchus and preliminary study of the hydrodynamic behaviour through computational fluid dynamics

This study characterized the morphology, density and orientation of the dermal denticles along the body of a shortfin mako shark Isurus oxyrinchus and identified the hydrodynamic parameters of its body through a computational fluid‐dynamics model. The study showed a great variability in the morphology, size, shape, orientation and density of dermal denticles along the body of I. oxyrinchus. There was a significant higher density in dorsal and ventral areas of the body and their highest angular deviations were found in the lower part of the mouth and in the areas between the pre‐caudal pit and the second dorsal and pelvic fins. A detailed three‐dimensional geometry from a scanned body of a shark was carried out to evaluate the hydrodynamic properties such as drag coefficient, lift coefficient and superficial (skin) friction coefficient of the skin together with flow velocity field, according to different roughness coefficients simulating the effect of the dermal denticles. This preliminary approach contributed to detailed information of the denticle interactions. As the height of the denticles was increased, flow velocity and the effect of lift decreased whereas drag increased. The highest peaks of skin friction coefficient were observed around the pectoral fins.     

Postnatal development of hepatic xanthine oxidase in baby pigs and turnover of purine catabolites at reduced ambient temperature.

1. The activity of xanthine oxidase in liver samples of baby pigs up to 4 weeks of age was investigated. On the 3rd day of life the turnover of hypoxanthine and of uric acid were measured after intravenous injection of 3H- and 14C-labelled tracers into animals kept at normal (32 degrees C) and reduced (20 degrees C) ambient temperature. 2. Hepatic xanthine oxidase activity increased progressively from 2 to 28 days of age (r = 0.689; P < 0.001). The increase of Vmax and of KM within 3-4 weeks was about 4.5-fold. 3. In 3-day-old baby pigs kept at normal temperature, pool size and turnover was about 10-fold higher for hypoxanthine than for uric acid. 4. At reduced ambient temperature, the pool size of uric acid increased 3.9-fold (P < 0.01) and turnover 1.6-fold (P < 0.05). For hypoxanthine the increases were insignificant.     

Teeth outside the mouth: The evolution and development of shark denticles

Vertebrate skin appendages are incredibly diverse. This diversity, which includes structures such as scales, feathers, and hair, likely evolved from a shared anatomical placode, suggesting broad conservation of the early development of these organs. Some of the earliest known skin appendages are dentine and enamel-rich tooth-like structures, collectively known as odontodes. These appendages evolved over 450 million years ago. Elasmobranchs (sharks, skates, and rays) have retained these ancient skin appendages in the form of both dermal denticles (scales) and oral teeth. Despite our knowledge of denticle function in adult sharks, our understanding of their development and morphogenesis is less advanced. Even though denticles in sharks appear structurally similar to oral teeth, there has been limited data directly comparing the molecular development of these distinct elements. Here, we chart the development of denticles in the embryonic small-spotted catshark (Scyliorhinus canicula) and characterize the expression of conserved genes known to mediate dental development. We find that shark denticle development shares a vast gene expression signature with developing teeth. However, denticles have restricted regenerative potential, as they lack a sox2+ stem cell niche associated with the maintenance of a dental lamina, an essential requirement for continuous tooth replacement. We compare developing denticles to other skin appendages, including both sensory skin appendages and avian feathers. This reveals that denticles are not only tooth-like in structure, but that they also share an ancient developmental gene set that is likely common to all epidermal appendages.     

Placoderm fishes,pharyngeal denticles,and the vertebrate dentition

The correlation of the origin of teeth with jaws in vertebrate history has recently been challenged with an alternative to the canonical view of teeth deriving from separate skin denticles. This alternative proposes that organized denticle whorls on the pharyngeal (gill) arches in the fossil jawless fish Loganellia are precursors to tooth families developing from a dental lamina along the jaw, such as those occurring in sharks, acanthodians, and bony fishes. This not only indicates that homologs of tooth families were present, but also illustrates that they possessed the relevant developmental controls, prior to the evolution of jaws. However, in the Placodermi, a phylogenetically basal group of jawed fishes, the state of pharyngeal denticles is poorly known, tooth whorls are absent, and the presence of teeth homologous to those in extant jawed fishes (Chondrichthyes + Osteichthyes) is controversial. Thus, placoderms would seem to provide little evidence for the early evolution of dentitions, or of denticle whorls, or tooth families, at the base of the clade of jawed fishes. However, organized denticles do occur at the rear of the placoderm gill chamber, but are associated with the postbranchial lamina of the anterior trunkshield, assumed to be part of the dermal cover. Significantly, these denticles have a different organization and morphology relative to the external dermal trunkshield tubercles. We propose that they represent a denticulate part of the visceral skeleton, under the influence of pharyngeal patterning controls comparable to those for pharyngeal denticles in other jawed vertebrates and Loganellia.     

Origin and evolution of gnathostome dentitions: a question of teeth and pharyngeal denticles in placoderms

The fossil group Placodermi is the most phylogenetically basal of the clade of jawed vertebrates but lacks a marginal dentition comparable to that of the dentate Chondrichthyes, Acanthodii and Osteichthyes (crown-group Gnathostomata). The teeth of crown-group gnathostomes are part of an ordered dentition replaced from, and patterned by, a dental lamina, exemplified by the elasmobranch model. A dentition recognised by these criteria has been previously judged absent in placoderms, based on structural evidence such as absence of tooth whorls and typical vertebrate dentine. However, evidence for regulated tooth addition in a precise spatiotemporal order can be observed in placoderms, but significantly, only within the group Arthrodira. In these fossils, as in other jawed vertebrates with statodont, non-replacing dentitions, new teeth are added at the ends of rows below the bite, but in line with biting edges of the dentition. The pattern is different on each gnathal bone and probably arises from single odontogenic primordia on each, but tooth rows are arranged in a distinctive placoderm pattern. New teeth are made of regular dentine comparable to that of crown-gnathostomes, formed from a pulp cavity. This differs from semidentine previously described for placoderm gnathalia, a type present in the external dermal tubercles. The Arthrodira is a derived taxon within the Placodermi, hence origin of teeth in placoderms occurs late in the phylogeny and teeth are convergently derived, relative to those of other jawed vertebrates. More basal placoderm taxa adopted other strategies for providing biting surfaces and these vary substantially, but include addition of denticles to the growing gnathal plates, at the margins of pre-existing denticle patches. These alternative strategies and apparent absence of regular dentine have led to previous interpretations that teeth were entirely absent from the placoderm dentition. A consensus view emerged that a dentition, as developed within a dental lamina, is a synapomorphy characterising the clade of crown-group gnathostomes. Recent comparisons between sets of denticle whorls in the pharyngeal region of the jawless fish Loganellia scotica (Thelodonti) and those in sharks suggest homology of these denticle sets on gill arches. Although the placoderm pharyngeal region appears to lack denticles (placoderm gill arches are poorly known), the posterior wall of the pharyngeal cavity, formed by a bony flange termed the postbranchial lamina, is covered in rows of patterned denticle arrays. These arrays differ significantly, both in morphology and arrangement, from those of the denticles located externally on the head and trunkshield plates. Denticles in these arrays are homologous to denticles associated with the gill arches in other crown-gnathostomes, with pattern similarities for order and position of pharyngeal denticles. From their location in the pharynx these are inferred to be under the influence of a cell lineage from endoderm, rather than ectoderm. Tooth sets and tooth whorls in crown-group gnathostomes are suggested to derive from the pharyngeal denticle whorls, at least in sharks, with the patterning mechanisms co-opted to the oral cavity. A comparable co-option is suggested for the Placodermi.     

A new species of <Emphasis Type="Italic">Trichodina</Emphasis> Ehrenberg, 1830 (Ciliophora: Trichodinidae) infecting farmed <Emphasis Type="Italic">Clarias gariepinus</Emphasis> (Burchell) (Siluriformes: Clariidae) in Cuba

A new species of Trichodina Ehrenberg, 1830 collected from the skin and fins of farmed North African catfish Clarias gariepinus (Burchell) fingerlings, is described. The new species can be distinguished from other trichodinids by the characteristics of the adhesive disc, especially by the great number of denticles. Trichodina merciae n. sp. is morphologically similar to T. renicola (Mueller, 1931) and T. marplatensis Martorelli, Marcotegui & Alda, 2008, in the number of denticles, but differs in the morphometric data, denticle morphology, environment and location. Trichodina merciae n. sp. has broad sickle-shaped blades and thin, straight rays, while T. marplatensis has broad club-shaped blades and wide S-shaped rays. Besides, denticle length, blade length, ray length, width of central part and denticle span of the new species are greater than T. marplatensis. However, the diameter of denticle ring and the diameter of the central area in T. marplatensis is larger than the ones in T. merciae n. sp. This is the first record of freshwater ectoparasite trichodinid with an average number of denticles greater than 50.