Twenty two percent (22/98) of intertidal fishes of 10 species captured in South Africa at Koppie Alleen, De Hoop Nature Reserve (south coast) and Mouille Point, Cape Town (west coast), harboured single or combined infections of haemogregarines, trypanosomes and an intraerythrocytic parasite resembling a Haemohormidium sp. The haemogregarines included the known species Haemogregarina (sensu lato) bigemina (Laveran et Mesnil, 1901) Siddall, 1995 and Haemogregarina (sensu lato) koppiensis Smit et Davies, 2001, while Haemogregarina (sensu lato) curvata sp. n. was observed in Clinus cottoides Valenciennes and Parablennius cornutus (L.) at Koppie Alleen. This last haemogregarine is characterised particularly by its distinctly curved gamonts. Also at Koppie Alleen, squash and histological preparations of 9/10 leeches, Zeylanicobdella arugamensis De Silva, 1963, taken from infected C. cottoides and P. cornutus contained developmental stages of H. curvata and/or trypanosomes, but these were absent from haematophagous gnathiid isopods (Gnathia africana Barnard, 1914) taken from infected fishes. It is suspected that Z. arugamensis transmits the haemogregarine and trypanosomes simultaneously between fishes, a double event unreported previously from the marine environment.
Blood films were examined from 154 wild and captive tortoises from four provinces of South Africa, including Gauteng, Kwazulu-Natal, North West and Western Cape. The five species of chelonians studied were Chersina angulata (Schweigger), Kinixys belliana belliana (Gray), K. lobatsiana Power, K. natalensis Hewitt, and Stigmochelys pardalis (Bell). Two species of haemogregarines, previously reported from Mozambique, were identified in blood films, namely Haemogregarina fitzsimonsi Dias, 1953 and Haemogregarina parvula Dias, 1953. Additional stages of development (trophozoites and probable meronts, merozoites and immature gamonts) in blood preparations from South Africa warranted the redescription of H. fitzsimonsi. A variety of hosts and broad host distribution range were observed for this haemogregarine, with all five species of tortoises parasitized, wild and captive, from all four provinces, in all seasons. In contrast, only two individuals of K. b. belliana and one S. pardalis, all three captive in Kwazulu-Natal, contained H. parvula with encapsulated stages resembling those of Hemolivia mauritanica (Sergent et Sergent, 1904). For H. fitzsimonsi, parasite prevalences, but not parasitaemias, were significantly higher in captive than wild S. pardalis; captive female S. pardalis also showed a significantly greater prevalence of infection than males, but younger, lighter hosts were not significantly more heavily parasitized than older, heavier individuals. The ticks, Amblyomma marmoreum Koch, 1844 and A. sylvaticum (De Geer, 1778), found attached to some tortoises, may prove to be definitive hosts for the two species of haemogregarines observed.
During 2001 and 2002, blood smears from 37 of 120 fishes belonging to 10 species captured in the Okavango Delta region of Botswana, were found to harbour trypanosomes. These trypanosomes displayed differing staining properties, were morphometrically variable, and ranged in total length from 29.5 to 80.8 µm. Mixed populations of the smaller and larger trypanosomes were found in most fish examined. Despite variations in size and appearance, these specimens are tentatively identified as Trypanosoma mukasai Hoare, 1932, likely adding another 9 new hosts to those known for this parasite. It is possible that Trypanosoma clariense Pienaar, 1962, described from Clarias gariepinus in South Africa, is also a junior synonym of T. mukasai.
Archived blood smears from 32 of 113 fishes in 18 families and 12 orders, trawled from deep North Atlantic waters off the Cape Verde Islands in 1999 and over the Porcupine Seabight in 2001 were found to harbour haematozoans. These included four species of haemogregarines (Adeleorina, Haemogregarinidae) and a species of trypanosome (Trypanosomatina, Trypanosomatidae) located in Porcupine Seabight fishes. Also present were Haemohormidium-like structures of uncertain status found in samples from this location and from the Cape Verde Islands. Although material was limited, two of the haemogregarines were provisionally named Desseria harriottae sp. n. from Harriotta raleighana Goode et Bean (Chimaeriformes, Rhinochimaeridae), and Haemogregarina bathysauri sp. n. from Bathysaurus ferox Günther (Aulopiformes, Bathysauridae). The two remaining haemogregarines were identified as Desseria marshalllairdi (Khan, Threlfall et Whitty, 1992) from Halosauropsis macrochir (Günther) (Notacanthiformes, Halosauridae), and Haemogregarina michaeljohnstoni (Davies et Merrett, 2000) from Cataetyx laticeps Koefoed (Ophidiformes, Bythitidae). The name H. michaeljohnstoni was proposed to replace Haemogregarina johnstoni Davies et Merrett, 2000 from C. laticeps and to avoid confusion with Hepatozoon johnstoni (Mackerras, 1961) Smith, 1996 from varanid lizards, originally named Haemogregarina johnstoni Mackerras, 1961. The trypanosome formed a mixed parasitaemia with D. harriottae in H. raleighana and was provisionally named Trypanosoma harriottae sp. n. No blood parasites had been described previously from cartilaginous fishes of the Holocephali, making the finds in H. raleighana unique. Haemohormidium-like structures were located in erythrocytes in one fish, Coryphaenoides armatus (Hector), among the Cape Verde Islands samples and in 12 species of fishes from the Porcupine Seabight; all these hosts were bony fishes. Finally, the haemogregarine species listed in the genus Desseria Siddall, 1995 were reassessed. Of the original list of 41 species, 30 were retained and 5 species added, including D. harriottae, so that the genus now contains 35 species.
This paper reviews past, current and likely future research on the fish haemogregarine, Haemogregarina bigemina Laveran et Mesnil, 1901. Recorded from 96 species of fishes, across 70 genera and 34 families, this broad distribution for H. bigemina is questioned. In its type hosts and other fishes, the parasite undergoes intraerythrocytic binary fission, finally forming mature paired gamonts. An intraleukocytic phase is also reported, but not from the type hosts. This paper asks whether stages from the white cell series are truly H. bigemina. A future aim should be to compare the molecular constitution of so-called H. bigemina from a number of locations to determine whether all represent the same species. The transmission of H. bigemina between fishes is also considered. Past studies show that young fish acquire the haemogregarine when close to metamorphosis, but vertical and faecal-oral transmission seem unlikely. Some fish haemogregarines are leech-transmitted, but where fish populations with H. bigemina have been studied, these annelids are largely absent. However, haematophagous larval gnathiid isopods occur on such fishes and may be readily eaten by them. Sequential squashes of gnathiids from fishes with H. bigemina have demonstrated development of the haemogregarine in these isopods. Examination of histological sections through gnathiids is now underway to determine the precise development sites of the haemogregarine, particularly whether merozoites finally invade the salivary glands. To assist in this procedure and to clarify the internal anatomy of gnathiids, 3D visualisation of stacked, serial histological sections is being undertaken. Biological transmission experiments should follow these processes.
Flounder, Paralichthys orbignyanus (Valenciennes), were captured in polluted and non-polluted sites within the Patos Lagoon Estuary, southern Brazil, over four seasons. Blood films showed a high prevalence of infection with a haemogregarine, or mixed parasitaemias of this and an organism resembling Haemohormidium terraenovae So, 1972. Haemogregarine gamont stages conformed to existing descriptions of Desseria platessae (Lebailly, 1904) Siddall, 1995 from flatfishes, but intraerythrocytic division of meronts was observed, leading to the recommendation for nomenclatural correction, placing the haemogregarine in the genus Haemogregarina (sensu lato) Danilewsky, 1885. Statistical analyses suggested that although sample sizes were small, infections with meront stages, immature and mature gamonts were all influenced by site, and possibly therefore, by pollution. Season also appeared to determine likelihood of infection with meronts and immature gamonts, but not mature gamonts, while adult fish gender apparently affected infection with immature and mature gamonts, but not meronts. The H. terraenovae-like organism exhibited unusual extracellular forms and did not match closely with the type description of H. terraenovae; precise identification was therefore difficult. Data analyses suggested that parasitism by this organism was influenced by site and fish gender, since females and males from non-polluted water were infected, but only females from the polluted site. Season was also important and significantly more adult fish of both sexes were infected with this parasite in the Brazilian summer and autumn, compared with winter and spring. Finally, these appeared to be the first observations of Haemogregarina platessae, and possibly H. terraenovae, from the southern hemisphere.
Haemogregarina bigemina Laveran et Mesnil, 1901 was examined in marine fishes and the gnathiid isopod, Gnathia africana Barnard, 1914 in South Africa. Its development in fishes was similar to that described previously for this species. Gnathiids taken from fishes with H. bigemina, and prepared sequentially over 28 days post feeding (d.p.f.), contained stages of syzygy, immature and mature oocysts, sporozoites and merozoites of at least three types. Sporozoites, often five in number, formed from each oocyst from 9 d.p.f. First-generation merozoites appeared in small numbers at 11 d.p.f., arising from small, rounded meronts. Mature, second-generation merozoites appeared in large clusters within gut tissue at 18 d.p.f. They were presumed to arise from fan-shaped meronts, first observed at 11 d.p.f. Third-generation merozoites were the shortest, and resulted from binary fission of meronts, derived from second-generation merozoites. Gnathiids taken from sponges within rock pools contained only gamonts and immature oocysts. It is concluded that the development of H. bigemina in its arthropod host illustrates an affinity with Hemolivia and one species of Hepatozoon. However, the absence of sporokinetes and sporocysts also distances it from these genera, and from Karyolysus. Furthermore, H. bigemina produces fewer sporozoites than Cyrilia and Desseria, although, as in Desseria, Haemogregarina (sensu stricto) and Babesiosoma, post-sporogonic production of merozoites occurs in the invertebrate host. The presence of intraerythrocytic binary fission in its fish host means that H. bigemina is not a Desseria. Overall it most closely resembles Haemogregarina (sensu stricto) in its development, although the match is not exact.