Politis, G.P.: Internal Parasites of a Sanddab Page 2 INTRODUCTION Parasitologists have been aware for many decades that fishes serve as hosts for a variety of helminth and protozoan parasites. In 1965, Mackenzie and Gibson reported a diverse parasite community ex¬ isting in the flat fishes flounder and plaice from the coast of England. Their study is probably the only detailed study ever published on para¬ site distribution and incidence in flat fish, and it is safe to say that there have been no published studies on parasites of the flat fish Citharichthys sordidus, the Pacific Sanddab. This paper describes five prevalent internal parasites of Citharichthys sordidus, and reports on the incidences of infection, average parasite loads per sanddab, and distributions of parasites within the host and host population. MATERIALS AND METHODS Sanddabs were collected by bottom fishing in Monterey Bay, nearly one mile northwest of Lovers Point, during the month of May. The study began with complete examinations of five sanddabs. The fishes were killed just prior to examination, by slicing the spinal cord immediately posterior to the operculum, and all the internal organs were inspected according to methods described by Amlacher (1970). Four parasites were found during this initial survey, and the study was narrowed to these more prevalent parasites. The weight, length, and sex were recorded prior to the inspection of each fish. The visceral cavity (Figure 1) was inspected with a di¬ Politis, G.P.: Internal Parasites of a Sanddab Page 3 secting microscope before any organs were removed. Physiological saline (0.9% Nacl) was periodically applied to the surface of organs during the examination. Gall bladder fluid was extracted with a sy¬ ringe and examined microscopically. The stomach and esophagus were re¬ moved, cut open, and viewed with a disecting microscope. The light source was put below the tissue, and forceps were used to spread the folds of the wall to locate hidden helminths. The intestine and py¬ loric caeca were examined in the same manner as the stomach and esoph¬ agus. The intestinal wall was inspected by cutting a small piece, rinsing it in distilled water, and flattening it for microscopic exam¬ ination. The ability of Trypanorhynchid larvae to survive an anaerobic en¬ vironment was evaluated by placing them in a stoppered test tube of deoxygenated (nitrogenated) physiological saline. The experiment was run at a temperature of 220 and a pH of 6. An attempt to induce Trypanorhynchid larvae to develop to their adult stage was done by putting the larvae in physiological saline solutions of pHs equal to 2, 3, and 4. Solutions were prepared by mix¬ ing dilute HCI with physiological saline, and using a pH meter to ob¬ tain the desired pHs. Viability of trypanorhynchid larvae was determined by preparing a slide and examining the scolex region for motion. When no motion was observed, gradually increasing pressure was applied to the coverslip until either motion was observed or the larva exploded. Politis, G.P.: Internal Parasites of a Sanddab Page 4 RESULTS The following data has been tabulated by assuming that parasite loads are independent of sanddab size and sex. The basis for this assumption was that Mann Whitney U-tests indicated no coorelation ex¬ ists between host sex and parasite loads, and scatter plots showed no coorelation between host size and parasite load. Fifteen out of fifteen gallbladders examined were infected by Ceratomyxa hopkinsi, a myxosporidean of the family Ceratomyxidae. Identification was made according to Kudo (1966). Ceratomyxa spores ranged in length from approximately 25um to 50gm, averaging about 40m. Mature digenetic trematodes of the family Hemiuridae (identifica¬ tion was according to Schell (1970)) were present in either the stomach or esophagus in 49% of the sanddabs. Figure 2 shows the hermaphroditic sac which is chacteristic of the family Hemiuridae. Hemiurids ranged from 1.5mm to 2.8mm, averaging 2.4mm (n=5), and had a mean load of 2.0 per sanddab. The concentration of hemiurids was greatest in the anter¬ ior end of the stomach; 21% were found on the esophagus, and 54% were found on the anterior half of the stomach (Table 1). Assuming that sampling of sanddabs was random, a Xø test determined that there was a significant difference between the number of trematodes found on the anterior half of the stomach and the number found on the posterior half (p-.05). Nematodes of the family Anisakidae were abundent in the pyloric caeca and intestine. The muscular pharynx and intestinal caecum were identifying characteristics, according to Hyman (1951). Six of 7 Politis, G.P.: Internal Parasites of a Sanddab Page 5 sanddabs were infected inside the intestine, with a mean of 3.6 Anis¬ akidae. Four of 7 were infected in the pyloric caeca, with a mean load of 1.6. The length of Anisakidae from inside the digestive tract ranged from 2mm to 8mm, with a mean of 5.4mm. Larval nematodes were found encysted in the walls of both the py¬ loric caecae and intestines in 6 out of 6 sanddabs. They were not encapsulated and were approximately .5mm in length. Anisakidae were also found in the body cavity, and occured with an incidence of 90%. They ranged from 5mm to 30mm in length, with a mean of 12.9nm. Of 57 anisakids from the visceral cavity, 30% were found attached to either the pyloric caeca, or it's adjacent mesentaries, and 30% were found on the mesentary in the cavity below the liver. On one occasion 7 metacercaria of the digenetic trematode Acantho¬ colpidae Stephanostomum sp. were found, and were attached to mesentary between the intestine and spleen. Plerocercoid larva of a cestode from the order Trypanorhyncha were abundant in the visceral cavity. These encapsulated larvae had a fully developed scolex, equipped with four spiny proboscides. The scolex region was invaginated in the receptaculum of the blastocyst (Figure 3) but could be extruded. Trypanorhynchids ranged from Imm to Amm in length, with a mean of 1.9mm (n-10), and had a mean load of 10 per sanddab. A concentration of trypanorhynchids were located in the area of the spleen and pyloric caeca; 28% were on the mesentaries adjacent to the spleen, and 28% were on the pyloric caeca or it's Politis, G.P.: Internal Parasites of a Sanddab Page 6 adjacent mesentaries. Trypanorhynchids were typically found in clumps, and often were in close proximity with nematodes; 11% had nematodes coiled around them. A Poisson distribution was calculated to determine what would be the expected number of sanddabs with various trypanorhynchid loads, pro¬ vided each infection was independent of all others (Figure 4). A X test value showed that the observed values were significantly different from the expected values (p is less than .01), indicating some type of dependence of infections upon one another. Five trypanorhynchid larvae were put into solutions of pH equal to 2, 3, and 4, to try to induce development. All five larvae in a pH of 2 had died by eight hours, while the larvae in pHs of 3 and 4 were still alive after 10 days. None of the larvae showed any sign of a morphological or behavioral change. Also, no change was observed in larvae that had been living for six weeks in a solution with a pH of 6. DISCUSSION The myxosporidean Ceratomyxa hopkinsi was consistently present in the gall bladder of Citharichthys sordidus. Jameson (1929) reported that Ceratomyxa hopkinsi was present in the gall bladder of Citharich¬ thys xanthostigma, but this is the first report of its occurence in Citharichthys sordidus. The significantly greater (p-.05) concentration of trematodes in the anterior region of the stomach is hard to explain. Mackenzie and Politis, G.P.: Internal Parasites of a Sanddab Page 7 Gibson (1970) reported that osmotic pressure of the gut fluid decreases down the length of the gut. If there is a difference in osmotic pressure within the stomach, the trematodes positioned anteriorly would be under greater osmotic stress and less likely to survive there. Perhaps the most interesting thing about the Hemiuridae data is the strange occur¬ ence of twenty-four hemiurids on a single sanddab. The life cycle of a digenetic trematode involves intermediate hosts and so we know the large number of hemiurids was not a result of reproduction on behalf of hemiurids already in the stomach. Possibly, metacercaria of this hemiurid are clumped within the population of the intermediate host on which sanddabs prey, or intermediate hosts are clumped in areas along the bottom. Taxonomy was a problem with the Anisakidae nematodes; the number of species of Anisakidae present in the sanddab was not determined. How¬ ever, all the Anisakidae found should be of the genera Contracaecum or Porocaecum, since Hyman (1951) reports that species of those two genera are the only anisakids found in marine fish. The occurence of larger nematodes in the visceral cavity andishorter ones in the digestive tract, indicates that the genus Contracaecum is probably present. According to Hyman, the typical life cycle of Contracaecum begins with eggs being passed into the water along with feces of the definative host. These eggs develop into an infective juvenile which is eaten by an intermedi¬ ate host (usually a fish). The juvenile develops in the host's intest¬ ine, and as it gets larger it bores through the intestine and inhabits body spaces. The intermediate host is eaten by the definative host Politis, G.P.: Internal Parasites of a Sanddab Page 8 and the life cycle is complete. Perhaps the sanddab's anasakids are immature Contracaecum with the sanddab serving as their intermediate host. Mackenzie and Gibson (1970) reported that Contracaecum aduncum was present in the flounder Platichthys flesus and in the plaice Pleuro¬ nectes platessa, taken from the coast of England. Contrary to Hyman, they state that mature Contracaecum were in the digestive tract and lar- vae were in the visceral cavity. The infections of trypanorhynchid larvae appear to be dependent on one another. Figure 4 shows that there is a disproportionately high number of fish with both few and many trypanorhynchids, implying that the distribution of larvae within the sanddab population is clumped. As with the trematodes, clumping is probably due either to clumping of larvae within the population of the previous intermediate host or to clumping of the previous host. The complete life cycle is not known for any marine trypanorhynchid, according to Hyman (1951). Rays and sharks are known to be definitive hosts, but it is not known how the larvae get to an intermediate host. Furthermore, how larvae get into the visceral cavity from inside the di¬ gestive tract is unknown. All trypanorhynchids found, including those encysted in the stomach wall and those inside the intestine, were encap¬ sulated and could not be using proboscides to bore. Interestingly, the trypanorhynchids found loose inside the intestine did seem more active than those in the visceral cavity; movement of the whole body was ob¬ served in the former while the latter were always motionless. Trypano¬ rhynchids in the visceral cavity may have entered a state of torpor. Politis, G.P.: Internal Parasites of a Sanddab Page 9 The fact that trypanorhynchids survive for over a week without oxygen indicates that they use a very small amount of oxygen or have an extraordinary capability to go into oxygen debt. Their ability to live for over six weeks in an anaérobic environment implies that their respiration rate is extremely low. Probably they enter a state of tor¬ por, and wait for an external cue to resume development. What the cue might be is unclear, but it does not seem to be a low pH. Coexistence of anisakids and trypanorhynchids suggests that there may be a symbiotic relationship between them. On the other hand, these helminths could be found together soley because of a shared interest in some resource or enviromental condition. Three points about this study are worth mentioning. Firstly, time was limited, making sample sizes smaller than desirable. Consequently, the assumption that there is not a coorelation between parasite loads and host size or sex could be incorrect. Secondly, an analysis of statisti¬ cal significance of trypanorhynchid and anisakid distributions could not be done since surface areas of organs and mesentaries were not com¬ puted. Thirdly, the information in this paper is probably specific for the community of Citharichthys sordidus studied, and for the time of year the study was done. The parasites of Citharichthys sordidus offer several opportunities for future work. Some possibilities include: illucidating the life cycle of Ceratomyxa hopkinsi, determining reasons for the observed helminth distributions, investigating the life cycle of the trypanorhynchid ces¬ tode, and determining the damaging effects of the various helminths on Politis, G.P.: Internal Parasites in a Sanddab Page 10 the host. SUMMARY 1. Five prevalent parasites were found during an investigation of 26 sanddabs. 2. Ceratomyxa hopkinsi, a myxosporidean, was present in 100% of the gall bladders examined. Trematodes of the family Hemiuridae were in the stomach and esophagus. A greater number were in the anterior stomach than the posterior. Nematodes of the family Anisakidae were in the intestine, pyloric caeca, and visceral cavity. Those in the visceral cavity were larger than those in the digestive tract. 5. Larval nematodes were in the wall of the intestine and pyloric caeca in 6 out of 6 examinations. 6. Cestode larvae of the order Trypanorhyncha were present in the visceral cavity. A concentration was noted in the area of the spleen and pyloric caeca. 7. Trypanorhynchid larvae do not randomly infect sanddabs; individual infections depend on one another. 8. Trypanorhynchid larvae can live in an anaerobic environment for at least 10 days. Politis, G.P.: Internal Parasites of a Sanddab Page 11 ACKNOWLEDGMENTS Thanks is due to several people who aided me during this study. I am grateful to: Dr. Robin Burnett for his statistical assistance and his comments on my sleeping habits; Dr. Norm Levine for his outgoing and modest nature; Dr Don Abbott for his enthusiastic attitude and his good ideas; Dr. Gary Wagenbach for his help editing this paper; Bill Magruder for his fishing talent; and Dragon Popoy for eating the sanddabs. Politis, G.P.: Internal Parasites of a Sanddab Page 12 LITERATURE CITED Amlacher, E.A., 1970. Textbook of Fish Diseases. T.F.H. Publications, Neptune City, New Jersey, pp. 25-26. Hyman, L.H., 1951. The Invertebrates, Vol. 3. McGraw-Hill, New York, New Vork. pp. 321-328. Jameson, A.P., 1929. Myxosporidia from California fishes. J. Para¬ sitology, 16:59. Kudo, R.R., 1966. Protozoology. Charles C. Thomas, Springfield, Illinois. pp. 774-806. Mackenzie, K. and Gibson D., 1970. Ecological Studies of Some Parasites of Plaice, Pleuronectes Platessa (L.) and Flounder, Platichthys Flesus (L.). In Taylor A.E. and R. Muller (Eds.), Aspects of Fish Parasitology. Sympos. British Soc. Parasitology, 8:1-42. Schell, S.C., 1970. How To Know The Trematodes. Wm. C. Brown Company, Dubuque, Iowa. Politis, G.P.; Internal Parasites of a Sanddab Page 13 Table Legends: Individual parasite distributions in Citharichthys sordidus. Table 1 Each value represents the percentage of the population of the parasite in the left hand column found on the organ listed above. Abbreviations are as follows: SP, splenic mesentary; P, pyloric caeca or adjacent mesentary; CL, mesentary in cavity under the liver; IM, intestinal mesentary; OS, on stomach wall; IS, inside stomach wall (encysted); E, in esophagus; AS, in anterior half of stomach; PS, in posterior half of stomach; IP, in pyloric caeca; I, in intestine; 0, other. 1 - 0 —. 30 8 S o o O o owoo o 8 oo ooo ooa- oaa o oopoo Politis, G.P.: Internal Parasites of a Sanddab Page 14 Figure Legends: Figure 1 Dorsal view of anatomical feature that are discussed in this paper. Figure 2 Ventral view of a flattened hemiurid tremitode, taken from the stomach. The distance from the center of the anterior sucker to the center of the ventral sucker represents approx. Imm. Figure 3 Flattened trypanorhynchid larvae from the visceral cavity. Figure 4 Representation of the expected number of sanddabs with a given trypanorhynchid load (according to a Poisson distribution) versus the observed number. The bars represent the observed number and the lines represent the expected. Groupings of trypanorhynchid loads were divided unequally to obtain expect- ed values of at least five so that a X test could be run. ESOPHAGUS LIVER BLADDER CAVITY UNDER LIVER RECTUM — PYLORIC STOMACH CARDIAC STOMACH SPLEEN PYLORIC CAECA Figure 1: Sanddab Anatomy Figure 2 Digenetic trematode from the stomach Figure 3 Trypanorhynchid larva from the visceral cavity Figure 4 Observed Expected 9—10 11—12 TRYPANORHYNCHID LOAD 13 Up 0 Appendix: Legend Parasites in individual Sanddabs. Also, Sanddab weight, length and sex. Abbreviations are as follows: Lg, length; Wt, weight; VA, visceral cavity anisakidae; DA, anisakidae from the digestive tract; LA, larval anisakidae; LT, larval trypanorhynchids; H, hemiuridae; CH, ceratomyxa hopkinsi. A "+" symbol means the para¬ site was present but too prevalent too count. A blank indicates that data was not collected. Appendix: (cm) Sanddab 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. Parasites in individual Sanddabs. 12 40 18 40 33 4 6 f LA + + 20 + O + 1 + 1 + 0 1 + O + 4 + O + 2 + 24 + 27 100 150 200 15