Revision of the Thelastomatoidea,Oxyurida of invertebrate hosts I. Thelastomatidae

Haemoproteus psittaci n. sp., a microsomal halteridial haemoproteid, is described from the African grey parrot Psittacus erithacus. This species differs from Haemoproteus handai, the only other haemoproteid described from the Psittaciformes, in that it is halteridial, not circumnuclear, is much smaller in all dimensions and has only 10–12 pigment granules compared to 20–24 noted in the latter species.     

The haemoproteids (Apicomplexa: Haemoproteidae) of the flowerpeckers of the avian family Dicaeidae (Passeriformes)

The haemoproteids of the flowerpecker family Dicaeidae are reviewed. Two new species, Haemoproteus dicaeus and H. nucleophilus, are described. Haemoproteus nucleophilus appears to be endemic in the dicaeids endemic to the New Guinea archipelago.     

Haemoproteus gabaldoni n. sp. (Apicomplexa: Haemoproteidae) from the Muscovy duck Cairina moschata (Aves: Anatidae)

Haemoproteus gabaldoni n. sp. is described from the Muscovy duck Cairina moschata from Caracas, Venezuela and is compared to H. greineri and H. nettionis which have been described previously from the Anatidae. The highly amoeboid outline, volutin granules and small number of pigment granules of H. gabaldoni serve to readily separate this species from the other two.     

ON THE FORAGING METHOD OF THE BANDED GYMNOGENE

Bennett, G. F., Earlé, R. A. &; Peirce, M. A. 1993. The Leucocytozoidae of South African birds: Musophagiformes, Cuculiformes and Piciformes. Ostrich 64:73-78.

The leucocytozoids of the avian orders Musophagiformes, Cuculiformes and Piciformes are reviewed. Leucocytozoon dinizi Tendeiro, 1947 of the Musophagidae is re-described and a neohapantotype designated. Leucocytozoon centropi Fantham, 1921 is re-described and L. coccyzus Coatney &; West, 1938 is synonymized with it and a neohapantotype designated. Leucocytozoon squamatus Nandi, 1986 of the Pi-cidae is re-described and compared with Leucocytozoon capitonis n. sp. of the piciform family Capitonidae.     

The haemoproteids of the avian families Eurylaimidae (broadbills) and Pittidae (pittas)

The haemoproteids of the avian suboscine families Eurylaimidae and Pittidae are reviewed. Haemoproteus eurylaimus n. sp. and H. pittae n. sp. are described.     

A STUDY OF THE SPECTACLED WEAVER

New species of haematozoa, namely Leucocytozoon ugwidi sp. nov. from the Cape Cormorant Phalacrocorax capensis and Haemoproteus skuae sp. nov. from the Subantarctic Skua Catharacta antarctica, are described. These are the first species to be recorded from the families Phalacrocoracidae and Stercorariidae, respectively.     

Blood Protozoa of Wild Turkeys in Florida*

From 1969 through 1972, 605 wild turkeys (Meleagris gallopavo Linnaeus) from 10 localities in Florida were examined for blood protozoans. The prevalence in turkeys more than a month old was 84% for Haemoproteus meleagridis Levine, 1961 and 72% for Leucocytozoon smithi Laveran and Lucet, 1905. Sixty-three percent of the birds were infected by both parasites. Infections were most prevalent in juveniles 8–12 months of age. No infection of H. meleagridis or L. smithi was found in 111 poults younger than 35 days. Prevalences were similar in both sexes. Haemoproteus meleagridis was more prevalent (87%) in the southern part of the state; L. smithi was more prevalent (75%) in the north. The prevalence of H. meleagridis did not change during the 4 year period, but L. smithi decreased markedly in 1971, a year of low rainfall. None of the 605 blood films was positive for Plasmodium, but 24 (75%) of 32 blood samples were found by subinoculation technics to be positive for a species of Plasmodium morphologically similar to P. durae Herman, 1941. No trypanosomes were seen on the 605 blood films or in bone marrow cultures (saline-neopeptone-blood) made from 11 turkeys.     

An annotated checklist of the valid avian species of Haemoproteus,Leucocytozoon (Apicomplexa: Haemosporida) and Hepatozoon (Apicomplexa: Haemogregarinidae)

The valid avian species of the apicomplexan blood parasite genera Haemoproteus, Hepatozoon and Leucocytozoon are arranged according to host family, assuming host familial specificity; salient points of the parasite morphology are recorded where appropriate.     

Phylogeny,Diversity, Distribution,and Host Specificity of Haemoproteus spp. (Apicomplexa: Haemosporida: Haemoproteidae) of Palaearctic Tortoises

A complex wide‐range study on the haemoproteid parasites of chelonians was carried out for the first time. Altogether, 811 samples from four tortoise species from an extensive area between western Morocco and eastern Afghanistan and between Romania and southern Syria were studied by a combination of microscopic and molecular‐genetic methods. Altogether 160 Haemoproteus‐positive samples were gathered in the area between central Anatolia and eastern Afghanistan. According to variability in the cytochrome b gene, two monophyletic evolutionary lineages were distinguished; by means of microscopic analysis it was revealed that they corresponded to two previously described species—Haemoproteus anatolicum and Haemoproteus caucasica. Their distribution areas overlap only in a narrow strip along the Zagros Mts. range in Iran. This fact suggests the involvement of two different vector species with separated distribution. Nevertheless, no vectors were confirmed. According to phylogenetic analyses, H. caucasica represented a sister group to H. anatolicum, and both of them were most closely related to H. pacayae and H. peltocephali, described from South American river turtles. Four unique haplotypes were revealed in the population of H. caucasica, compared with seven haplotypes in H. anatolicum. Furthermore, H. caucasica was detected in two tortoise species, Testudo graeca and Testudo horsfieldii, providing evidence that Haemoproteus is not strictly host‐specific to the tortoise host species.     

Haemoproteus in Wild Turkeys from the Coastal Bend of South Texas*

SYNOPSIS. Blood smears from 133 Rio Grande wild turkeys Meleagridis gallopavo intermedia live-trapped in the Welder Wildlife Refuge in south Texas, were examined for blood parasites. All of the 63 immature turkeys and 63% of the 70 adult turkeys were infected with Haemoproteus meleagridis. The infections were light, half the cases having less than 5 infected cells per 10,000 red blood cells. Infection with H. meleagridis caused significant enlargement of the host RBC. Other parasites specifically searched for but not found were Leucocytozoon, Trypanosoma, Plasmodium and microfilariae.     

An Electronmicroscopic Study of the Microgamete of Haemoproteus columbae Kruse*

SYNOPSIS. Microgametes of Haemoproteus columbae were studied by light microscopy and electron microscopy, using the single-stage replication technic. The filiform microgamete has a swelling or vesicle near the round end; the opposite end is asymmetrically tapered. The gamete is polarized, for it travels in sinuous movements with the round end (the putative anterior end) directed forward. There is no flagellum or undulating membrane and no mitochondrion. The vesicle may contain energy reserves for locomotion or lytic enzymes for penetration. Tubular material often is seen coming from the vesicle and anterior end and may be a kind of acrosomal filament.     

Some Blood Sporozoans from East African Reptiles*

SYNOPSIS. Six hundred and forty-one lizards belonging to 28 species, and 29 snakes belonging to 12 species from East Africa were examined for blood sporozoans. Eight species of lizards and 4 of snakes were positive. Tissue phases were seen in some infections; no proven arthropod vectors were found. Parasites are described from previously unreported hosts; new host records, and new stages in the life histories of previously named parasites are reported. The sporozoans belong to the genera Plasmodium, Haemoproteus, Haemogregarina or Hepatozoon, and probably Pirhemocyton. Reasons are given for the inadvisability of establishing new species of some of these on the basis of blood stages alone.     

Two new species of Haemoproteus Kruse, 1890 (Haemosporida,Haemoproteidae) from European birds,with emphasis on DNA barcoding for detection of haemosporidians in wildlife

Two new species of Haemoproteus Kruse, 1890 (Haemosporida, Haemoproteidae) are described: Haemoproteus (Parahaemoproteus) homovelans n. sp. from Grey-faced Woodpecker, Picus canus Gmelin, and Haemoproteus (Parahaemoproteus) concavocentralis n. sp. recorded in Hawfinch, Coccothraustes coccothraustes (Linnaeus), both sampled in Bulgaria. The morphology of the gametocytes and their host-cells are described and mitochondrial cytochrome b (cyt b) gene sequences are generated. Haemoproteus homovelans possesses circumnuclear gametocytes lacking volutin granules. This parasite is particularly similar to Haemoproteus velans Coatney & Roudabush, 1937 also possessing circumnuclear gametocytes that are, however, overfilled with volutin. Haemoproteus concavocentralis can be readily distinguished from all described avian haemoproteids due to the presence of an unfilled concave space between the central part of advanced gametocytes and erythrocyte nucleus. Bayesian phylogenetic analyses of 40 haemosporidian cyt b lineages showed close relationships of H. concavocentralis (hHAWF2) with a group of Haemoproteus spp. possessing gametocytes that are pale-stained with Giemsa. The lineage hPICAN02 of H. homovelans clustered with parasites infecting non-passerine birds. Phylogenetic analyses support the current subgeneric classification of the avian haemoproteids and suggest that cyt b lineage hPIPUB01 (GenBank EU254552) has been incorrectly assigned to Haemoproteus picae Coatney & Roudabush, 1937, a common parasite of corvid birds (Passeriformes). This study emphasises the importance of combining molecular techniques and light microscopy in the identification and field studies of avian haemosporidian parasites. Future development of barcodes for molecular identification of haemoproteids will allow better diagnostics of these infections, particularly in veterinary studies addressing insufficiently investigated tissue pathology caused by these parasites.     

Pre-Erythrocytic Development and Associated Host Responses to Haemoproteus meleagridis (Haemosporina: Haemoproteidae) in Experimentally Infected Domestic Turkeys12

ABSTRACT. Two generations of pre-erythrocytic schizogony occurred in skeletal and cardiac muscle of domestic turkeys infected with sporozoites of Haemoproteus meleagridis. First generation schizonts reached maturity approximately five days post-inoculation (DPI) and developed in capillary endothelial cells and myofibroblasts. The schizonts ranged from 12 to 20 μm in diameter and produced long (5–6 μm), slender merozoites. Early second generation schizonts were first detected in capillary endothelial cells between 5 and 8 DPI. They were cylindrical and ranged in size from 5 to 8 μm in diameter and up to 28 μm in length. Second generation schizonts which reached maturity by 17 DPI were surrounded by a thick, hyaline wall and were packed with numerous spherical merozoites less than 1 μm in diameter. Mature megaloschizonts were fusiform, ranged from 30 to 113 μm in diameter, and extended as much as 465 μm along the long axis of muscle fibers. Merozoites developed as buds from cytomeres that formed between 8 and 14 DPI. Infected turkeys developed a moderate to severe myositis within 5 DPI and were lame in one or both legs. The myositis was associated with the necrosis of scattered groups of muscle fibers. Muscle fibers surrounding mature megaloschizonts were swollen and hyaline. Megaloschizonts were surrounded occasionally by fibroblasts and infiltrates of mononuclear cells. The morphology and site of development of mature megaloschizonts of Haemoproteus meleagridis are contrasted with those of other avian haemosporidians.     

Fine Structure of Haemoproteus columbae Kruse During Macrogametogenesis and Fertilization*

When blood is withdrawn from a pigeon (Columba livia) infected with gametocytes of Haemoproteus columbae, differentiation of the gametes begins immediately. This study examines the formation of the macrogamete and its fertilization. The first visible signs of differentiation are the elongation of the nucleus along with the appearance of an intranuclear spindle and atypical centrioles. Then maturation bodies, the products of nuclear reduction, form in both erythrocytic macrogametocytes and macrogametocytes free of their host cells. Penetration of the macrogamete by the microgamete occurs rapidly. Their plasma membranes fuse, and the microgamete's nucleus, axonemes and cytoplasm enter the macrogamete. The nucleus of the microgamete expands and migrates to lie at an angle to the macrogamete nucleus. The 2 fuse across a small area. The nuclear envelope and the plasma membrane of the zygote are a mosaic of the membranes of the 2 gametes.     

Observations on the Fine Structure of the Schizonts of Haemoproteus columbae Kruse*

SYNOPSIS. The schizonts of Haemoproteus columbae resemble the exoerythrocytic schizonts of avian Plasmodium in their fine structure. Haemoproteus infects endothelial cells and grows several hundredfold in volume, destroying the cytoplasm and nucleus of the host cell. The schizont's plasma membrane is trilamellar with a dense outer lamella. Some schizonts have micropores in their plasma membranes, but there is no evidence for ingestion thru them. Instead, numerous vesicles and channels fill the host cell cytoplasm and give its plasma membrane and periparasitic vacuolar membrane the appearance of active pinocytosis. The parasite's membrane shows no sign of pinocytosis, indicating that it probably feeds by diffusion. The growing schizont has numerous mitochondria, nuclei, and ribosome-rich cytoplasm which contains electron-lucent vacuoles and clefts. The latter appear to be artifacts of fixation.     

Are chronic avian haemosporidian infections costly in wild birds?

One group of commonly found parasites in birds, for which fitness consequences and effects on life history traits have been much debated are Haemosporidian blood parasites. In a long term study population of great reed warblers Acrocephalus arundinaceus in Sweden, previous studies have shown that the Haemosporidian blood parasites are in their chronic phase during the breeding season and that the fitness of infected and non‐infected birds are similar. In the present study, we quantified parasite intensity (parasitemia) in 718 adults great reed warblers sampled between 1987 and 1998 for the three most common parasite species; Haemoproteus payevskyi (lineage GRW1), Plasmodium ashfordi (GRW2) and Plasmodium relictum (GRW4). We verified that the q‐PCR method is accurately quantifying Haemoproteus payevskyi (GRW1) as it was highly correlated with the number of parasites seen under microscope. Frequency of mixed infections with two lineages was significantly higher than expected based on the prevalence of each of the three parasite lineages. The mean level of parasitemia was significantly different for the three lineages and individual birds had repeatable parasitemia levels between years. Females tended to have a higher parasitemia than males for all three parasite lineages combined. Females with higher GRW1 parasitemia tended to arrive later in spring to their breeding sites. There was a negative correlation between parasitemia and number of fledged offspring for GRW1, and a tendency for a negative correlation between GRW2 parasitemia and the proportion of recruiting offspring. Overall our results demonstrate that chronic Haemosporidian infections can have slight but significant effects on host life history traits, and therefore may act as important selective agents in wild bird populations.     

Blood Parasites of Taiwan Birds*

SYNOPSIS. Blood films were examined from 1477 birds of Taiwan (193 species, 49 families). Haemoproteus Kruse was by far the commonest parasite, with Leucocytozoon Danilewski a not very close second. It is probable that some of the Haemoproteus infections represented new species, and 1 occurring in the Bamboo Partridge (Bambusicola thoracica sonorivox Gould) seemed characteristic enough to justify recognition as such; the name Haemoproteus bambusicolae sp. n. is proposed for this organism. Malaria was found in 77 birds, the greatest number of infections occurring in the Bamboo Partridge. Most of them were caused by Plasmodium juxtanucleare Versiani & Gomes, a pathogen of chickens, but a number were due to an undetermined species of Plasmodium. The Bamboo Partridge may be a reservoir host of the former. A few other identified species (P. rouxi Sergent & Sergent, P. hexamerium Huff, P. tenue Laveran & Mesnil) were seen, as well as some unidentified ones. Plasmodium tenue was seen in Garrulax canorus taewanus Swinhoe, a babbler: until now it was known only from the Pekin Robin (Leiothrix luteus Scopoli), also a babbler, in which we have found it extremely common. Sixty-four microfilarial infections were identified; they were especially frequent in the Button Quail (Turnix suscitator rostrata Swinhoe).     

A Comparative Study of Gametocyte Ultrastructure in Avian Haemosporidia*

The fine structure of gametocytes of 3 avian haemosporidian parasites Plasmodium gallinaceum, Haemoproteus columbae, and Leucocytozoon simondi has been studied and compared by electron microscopy. The gametocytes of all 3 species are bounded by a 3-layered limiting membrane system, possess a cytostome during some portion of their residence within host cells, and their sex can be distinguished by both nuclear and cytoplasmic characteristics. L. simondi differs most significantly from P. gallinaceum and H. columbae in possessing large intranuclear granules, mitochondria associated with pocket infoldings of the nuclear envelope near the atypical centriole complex and compartmentalization of the cytoplasm by segments of closely aligned unit membranes. Further, the cytostome of L. simondi does not appear to be a persistent structure as in the other 2 species and pigment is not present within food vacuoles. L. simondi also is capable of infecting a wider variety of host cells and within leukocytes produces striations of the host nucleus and an apparent spiral banding of the host cell surface. The comparison of P. gallinaceum, H. columbae, and L. simondi gametocytes by electron microscopy leads to the conclusion that Plasmodium and Haemoproteus are more closely related to each other than either of them is on Leucocytozoon. The terminology used to describe certain organelles within the gametocyte's cytoplasm has been reexamined and the relationship of the nucleolus to parasite maturation also is described.     

Fine Structure of Differentiating Oocysts and Mature Sporozoites of Haemoproteus metchnikovi in its Intermediate Host Chrysops callidus*

The fine structure of the sporogonic stages of Haemoproteus metchnikovi has been investigated by electron microscopy. Young oocysts are found beneath the basement membrane of midgut epithelial cells. These eventually protrude outward into the haemocoel space and are surrounded by a distinct oocyst capsule. Sporozoite formation begins with a subcapsular vacuolation. Evagination of the oocyst cytoplasm occurs in regions of membrane thickenings and 100–200 sporozoites are formed about a single sporoblastoid body. Remnants of the ookinete pellicle can be observed in maturing oocysts and always are found in the residual body. The fine structure of the mature sporozoite is essentially similar to that which has been described for other haemosporidia and a spherical body is described in association with the mitochondrion of the sporozoite. The sporogonic stages of H. metchnikovi have features common to the sporogonic stages of Plasmodium and Leucocytozoon that are not held in common by the latter 2 genera, including pattern of sporozoite formation and number of sporozoites formed, the presence of a cytostome and of “crystalloid” in the sporozoite.