Apicomplexa Secondary article

William C Marquardt, Colorado State University, Fort Collins, Colorado, USA Article Contents
CA Speer, Montana State University, Bozeman, Montana, USA . Introduction
. Characterization
. Place in Overall Taxonomic Scheme
The phylum Apicomplexa includes several thousand parasitic protists that cause major
. Phylogenetic and Evolutionary Considerations
diseases of mankind and of domestic and companion animals. Among the diseases that
. Host–Parasite Interactions
they cause are malaria, coccidiosis, toxoplasmosis, sarcocystosis, cyclosporosis,
. Major Taxa and Important Species
cryptosporidiosis and piroplasmosis.

Introduction (Figure 2), often referred to as a micropore. All species are

The Apicomplexa comprises a phylum of protists that have parasitic.
structures referred to as the apical complex and in which Schizogony is an asexual process of multiplication in
the life cycles have both asexual and sexual phases. All which multiple nuclear replications take place without
members are parasitic. Included in the phylum are several
thousand described species that parasitize members of all
of the major animal phyla. They are parasites of common
invertebrate phyla such as molluscs, arthropods and
annelids as well as of some lesser-known phyla. Apicom-
plexans are parasites of members of all the classes of the
phylum Chordata including sea squirts or tunicates, fish,
amphibia, reptiles, birds and mammals.
Members of the Apicomplexa cause some of the most
devastating diseases of mankind and cause enormous
economic losses in domesticated animals and poultry. The
most prominent apicomplexans are Plasmodium spp.,
which cause malaria in humans; even after one hundred
years of concerted control programmes, malaria still takes
more than 2 million human lives each year. Coccidiosis is a
complex of intestinal diseases of livestock and poultry that
are common wherever animals are raised for food and
wool. Global losses to coccidiosis reach into many millions
of dollars (US) annually. Poultry cannot be raised in
current high concentration methods without the contin-
uous use of drugs to prevent the development of the
causative agents of coccidiosis. Piroplasmoses are tick-
borne diseases that affect livestock on all continents and
may be the single most important group of infectious
agents of grazing animals.

Characterization
Members of the phylum have one or more elements of the
apical complex (Figure 1) at some stage of the life cycle:
polar ring, rhoptries, micronemes and conoid. Sexuality is Figure 1 Structure of apicomplexans as revealed by electron microscopy.
by syngamy. Multiplication is by schizogony or endodyo- Upper left: a metrocyte, or mother cell, in which the elements of the apical
geny; 9 1 2 flagellar form is present in some gametes; complex are minimally expressed. Upper right: a zoite in which all of the
elements of the apical complex are seen. Lower: A detailed view of the
locomotion is by body flexion or gliding; subpellicular conoid and associated elements such as the polar rings and the
tubules are present in most species. Feeding is by subpellicular tubules which extend about two-thirds of the length of the
osmotrophy or phagotrophy, the latter by a cytostome zoite. (Courtesy of C. A. Speer.)

ENCYCLOPEDIA OF LIFE SCIENCES © 2001, John Wiley & Sons, Ltd. www.els.net 1
 Apicomplexa

cytokinesis until all the progeny have formed. Then, organisms such as Plasmodium spp. Merogony is also a
cytokinesis occurs simultaneously forming all of the schizogonic process and the main one by which the
progeny (Figure 3). Endodyogeny is a process in which apicomplexans multiply within a host. In malaria, this
two progeny form within a parental cell (Figure 4); when the process occurs first in the liver and then in the red blood
offspring are fully formed, the parental cell breaks down, cells. Gamogony results in the formation of gametocytes,
releasing them. which undergo syngamy to form a new individual. In most
Apicomplexans have three stages in their life cycles: instances, there are distinctly male and female gametes, but
in many of the gregarines, isogametes are formed.
Sporogony

Place in Overall Taxonomic Scheme

Gamogony Merogony Protistan (protozoan and protophytan) classification has
undergone and continues to undergo radical changes.
Note that the cycle runs in only one direction. The Despite what has occurred in other taxa, the phylum
organism is genetically programmed for one stage to Apicomplexa remains a discrete group. The organelles that
follow the other. comprise the apical complex as well as the life cycles and
Eimeria tenella of the chicken is a typical apicomplexan mode of division are quite consistent within the group. The
that has its stages in the intestinal tract of its host (Figure 5). exception is the class Perkinsea whose inclusion within the
Sporogony takes place outside of the host when oocysts are phylum remains problematic. Only three species have been
shed with the faeces. When an infective oocyst (Figure 6) is described and they have been included in the phylum
eaten, merogony takes place in the cells of the large Apicomplexa based on a structure that appears to be a
intestine. After three merogonies, gamogony takes place partial conoid but molecular studies will be needed to
with the formation of macro- and microgamonts. Upon clarify the status of this enigmatic group.
syngamy, the cycle is completed with the formation of
oocysts.
Sporogony is a schizogonic process in which many
sporozoites are the product and which usually takes place Phylogenetic and Evolutionary
within an oocyst (Figure 6). The sporozoite is the infective Considerations
stage in homoxenous parasites and in heteroxenous
In classification and in evolutionary studies, one tries to
determine which group serves as an ancestor for another.
In the case of protists generally, reaching such conclusions
was rather difficult until the development of electron
microscopy and molecular methods. The evidence now
suggests that Apicomplexa arose from a mostly marine
taxon commonly referred to as the dinoflagellates. Most of
its members are free-living, but some are symbiotic in
metazoa. The conclusion that the apicomplexans are
derived from the dinoflagellates is based on (1) the types
of hosts infected, (2) the mode of division (schizogony) in
some members, (3) cyst formation, and (4) the nucleic acid
sequence of the small subunit rRNA gene. A peculiar
multiple-membrane bound, DNA-containing, plastid-like
organelle (originally called a Golgi adjunct) has been
described in several apicomplexans. Phylogenetic analysis
of the plastid gene (called tufA) of Toxoplasma gondii has
placed its organellar genome with cyanobacteria and
plastids, especially green algal plastids. An intimate
association between the plastid and the nuclear spindle
has also been found in some apicomplexans, indicating
that the plastid may play a role in nuclear division (Speer
Figure 2 The cytostome or micropore of Toxoplasma gondii as seen in and Dubey, 1999).
transmission electron microscopy. The cytostome ingests food provided by Within the phylum Apicomplexa itself, the following is
the parasite’s host cell. (Courtesy of C. A. Speer.) generally agreed upon as the evolutionary sequence for the

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Figure 3 A meront of Sarcocystis sp. in a hepatocyte of a mouse as seen in transmission electron microscopy. (A) Merozoites that have completed budding
from (B) a centrally located residual body. (Courtesy of C. A. Speer.)

classes (excluding Perkinsea). Probably first were gregar- Haemosporea and Piroplasmea transmission haemato-
ines, or something like them. These organisms infect phagous arthropods inject the organisms.
invertebrates and most are coelozoic although some
groups are cytozoic. The long evolutionary history of the
apicomplexans can be inferred from the fact that members
of the genus Eimeria parasitize tunicates or sea squirts,
marine chordates from which fishes began to diverge Host–Parasite Interactions
perhaps 500 million years ago. Eimeria spp. infect the
intestines of all classes of vertebrates: fish, amphibia, Nearly all apicomplexans are cytozoic intracellular para-
reptiles, birds and mammals. Eimeria spp., with few sites. The gregarines usually have a holdfast or epimerite by
exceptions show a high degree of host specificity, usually which they attach to cells; as the organisms grow, the
infecting only a few closely related species of hosts and trophozoites become large enough to see with the naked
suggesting a long coevolution. eye and extend into the cavity of the organ. In Plasmodium
The shift to a two-host life cycle, such as occurs spp., which cause malaria in humans, the stages in the
facultatively in Toxoplasma gondii, probably occurred mosquito are extracellular. The intracellular mode of life is
when a paratenic host was inserted into the life cycle. Two often said to protect the organism from the immune
hosts eventually became obligatory, as in Sarcocystis and response of the host, but the long association of
Neospora. Transmission by haematophagous arthropods apicomplexans and coevolution with their hosts also
(or other invertebrates) is the final step in the evolutionary suggests that they parasitize the metabolic reactions of
sequence. In the Adeleina, the arthropod, often a their host cells. The striking enlargement of the nuclei and
mosquito, is eaten in order to infect the vertebrate. In nucleoli in host cells seen in certain coccidial infections

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Figure 4 Endodyogeny in zoites of Neospora caninum, an economically important parasite of domestic animals. (A) A zoite not undergoing
endodyogeny. (B) Partially formed progeny cells in which the nucleus is being divided into two progeny cells. (C) Two progeny cells completely formed and
probably ready to be released. (Courtesy of C. A. Speer.)

suggests that the transcriptional activity of the host cells is Sarcocystis, Toxoplasma, Isospora, Hammondia and
increased, perhaps selectively. others. In vitro cultivation of apicomplexans has led to
information on antigens, parasite development, biochem-
istry, host–parasite interactions, host cell invasion and
In vitro cultivation molecular biology.
A major advance concerning the cultivation of apicom-
Cultivation of an infectious agent outside of its host allows
plexans was made in 1976 by Trager and Jensen who
the organism to be characterized and its mode of
succeeded in continuous cultivation of meronts of Plas-
development can readily be studied. Advances in cultivat-
modium falciparum, the organism that causes malignant
ing bacteria in the nineteenth century brought about
tertian malaria in humans. This discovery facilitated
enormous advances in knowledge of infectious diseases
research in malaria immunology and biochemistry and is
and in their control. The earliest attempts to culture
recognized as initiating the era of molecular biological
apicomplexan parasites were made early in the twentieth
research on malaria.
century and involved Plasmodium spp., Sarcocystis tenella,
In a few instances, the complete life cycles of apicom-
Gastrocystis gilruthi, Toxoplasma gondii and Eimeria spp.
plexans have been obtained in vitro. In most cases, when
These attempts were unsuccessful, but somewhat later,
cultured cells are inoculated with sporozoites, apicomplex-
techniques for isolating and cultivating cells and tissues
ans develop only through the first-generation meronts or
from vertebrates were refined. By the late 1940s, major
they multiply exclusively by endodyogeny. Some apicom-
discoveries led to the development of reliable methods of
plexans are facultative or obligately heteroxenous para-
culturing vertebrate cells in vitro. Subsequently, techniques
sites (i.e. they require two hosts to complete their life
were developed that led to standardized procedures for the
cycles). Even in those that are monoxenous, the endogen-
cultivation of many types of cells, to the development of
ous stages often develop in different sites and in different
highly defined culture media, and to pathogen-free culture
types of cells within the host. For example, first-generation
media. Today, various culture media are available
meronts of Eimeria bovis develop in endothelial cells of the
commercially, as are some apicomplexan parasites.
central lacteals of the ileal portion of the intestine, whereas
Techniques have now been developed for the in vitro
second-generation meronts and gametocytes develop in
cultivation of many apicomplexans including species of
glandular epithelial cells in the large intestine and caecum.
Babesia, Plasmodium, Cryptosporidium, Eimeria, Theileria,

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Figure 5 The life cycle of Eimeria tenella of the chicken. This apicomplexan is typical of those coccidia that parasitize the intestinal tracts of their hosts.
Sporogony takes place outside of the host, and both merogony and gamogony occur within cells of the intestine. (From Marquardt et al., 1999.)

Cultures inoculated with sporozoites of E. bovis develop Neospora caninum in vitro. Meronts and gametocytes of
only to first-generation meronts, but merozoites obtained these species have not been cultured in vitro, indicating that
from the intestinal tract of infected calves develop to in these systems the appropriate cell types, cultural
gametocytes and oocysts in cultured cells. Further parasite conditions or components of the medium are lacking.
development probably involves certain environmental Research has shown that lowering the pH of the culture
conditions that are required for the stimulation of medium or adding parasite-specific antibodies to the
developmentally regulated genes, enabling the parasite to culture medium induces tachyzoites of T. gondii, Ham-
proceed to the next developmental phase. mondia spp. and N. caninum to form cysts containing
The complete endogenous cycles of Cryptosporidium bradyzoites. Techniques have also been developed for the
parvum, Isospora suis and certain Eimeria spp. from isolation and cultivation of apicomplexans from tissues of
chickens have been obtained in vitro. Endodyogeny is the infected hosts. For example, apicomplexans such as T.
only form of asexual reproduction that occurs with gondii, N. hughesi and Sarcocystis neurona (the latter
Toxoplasma gondii, Besnoitia spp., Hammondia spp. and

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a variety of cell types from a wide range of hosts, indicating

that the biochemical receptors involved in attachment and
penetration involve either the presence of a highly
conserved host cell receptor or a variety of different
receptors recognized by the parasite. In T. gondii,
tachyzoite attachment to host cells involves ubiquitous
receptors consisting of a parasite laminin–host cell laminin
receptor, lectins, and a glycosylated receptor, SAG1.
Attachment of tachyzoites to host cells can be inhibited
by antibodies and is saturable in peptide competition
assays. Tachyzoites made deficient in SAG1 bind signifi-
cantly less to host cells than do wild-type tachyzoites. In
addition, micronemes of T. gondii tachyzoites contain at
least three specific proteins (MIC1–3), some of which are
exocytosed during host cell penetration and may be
involved in host cell receptor recognition and attachment.
Proteins containing one or more copies of the type I
repeat of human platelet thrombospondin (TSP1) are
necessary for parasite motility and invasion of host cells.
Members of the vertebrate TSP family are adhesive
molecules involved in cell–cell and cell–matrix interactions
and blood coagulation. The presence of TSP1 repeats in
apicomplexans appears to enable these parasites to mimic
host cell molecules, such as TSP1, F-spondin, properdin
and C6-C9 of the complement cascade, and use their
natural ligands during movement over the surface of host
cells and during host cell penetration. Among the
Figure 6 An oocyst of Eimeria magna of the rabbit, Oryctolagus cuniculi, as Apicomplexa, proteins related to motility include all of
seen in interference contrast light microscopy. The oocyst is infective if the proteins released by sporozoites of Plasmodium species,
ingested by a rabbit. In the genus Eimeria, each oocyst has four sporocysts the thrombospondin-related adhesive protein (TRAP) of
each of which contains two sporozoites. (A) Oocyst wall. (B) Sporocyst. (C)
P. falciparum and other Plasmodium species, the micro-
Sporozoite. (Courtesy of C. A. Speer.)
nemal proteins of Eimeria maxima (Em100) and E. tenella
(Etp100), micronemal protein 2 (MIC2) of Toxoplasma
causes equine protistan myeloencephalitis) can be isolated
gondii and the TRAP of Cryptosporidium parvum. Based on
from brain and spinal cord tissue and grown in vitro.
studies with these apicomplexans, a molecular model has
been proposed for TRAP function in zoite motility. TRAP
Penetration of host cells is stored within micronemes at the sporozoite apical end
and is continuously translocated to the zoite plasmalem-
Zoites of many apicomplexans are highly motile, under- ma. TRAP binds to extracellular host cell ligands and acts
going flexing, gliding, and pivoting, and probing with their as a link with the parasite molecular motor. A network of
anterior ends. Direct microscopic observation has revealed actin–myosin lies in direct contact with the outer face of the
that host cell penetration is usually preceded by gliding zoite plasmalemma. Backward sliding of the myosin–
movements and probing of the host cell surface with the TRAP–ligand complexes along longitudinal actin fila-
parasite’s anterior end. Parasite motility and cell penetra- ments pushes the zoite body forward. This causes
tion are powered by an actin–myosin based motor in the accumulation at the posterior of the zoite of TRAP
parasite that is characterized by the rearward capping of complexes, which dissociate leaving a trail of TRAP.
surface membrane proteins that propels the parasite Apicomplexans have also been found to enter cells by
forward in a helical path. being phagocytosed or by purely mechanical mechanisms.
In general, the events involved in zoite attachment and For example, sporozoites of E. bovis that were completely
penetration of host cells include: (1) gliding of the zoite surrounded by host cell membranes and cytoplasm have
immediately before attachment and penetration; (2) been observed by transmission electron microscopy to
probing and attachment to the host cell with the conoid; penetrate other cells. Because sporozoites were still
(3) indentation of the host cell plasmalemma; (4) formation surrounded by an envelope containing host cell mem-
of a junction that moves posteriorly as the zoite penetrates branes, the requirement for contact between the tip of the
the host cell; and (5) partial exocytosis of micronemes, sporozoite and the receptor on the host cell plasmalemma
rhoptries, and dense granules. In vitro, zoites can penetrate may have been fulfilled, enabling the sporozoite to

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penetrate another cell. However, there would be no usually have elevated levels of tumour necrosis factor alpha
receptor contact on the plasmalemma of the host cell (TNF-Ã). Localized overproduction of TNF-Ã is con-
being penetrated by a host cell-enveloped sporozoite. sidered to be primarily responsible for fever and tissue
Therefore, it appears that in some instances host cells may lesions in severe falciparum malaria, but IgE also appears
be receptive to purely mechanical penetration by zoites to contribute to the pathogenicity.
without the need of host cell–parasite receptor contact. More than 60 years have been dedicated to searching for
Penetration of an apicomplexan into a cultured cell does a malaria vaccine, yet there is still no effective means of
not necessarily result in multiplication of the parasite. immunizing against the disease. Four species infect
After entry into host cells, the parasite and host cell must humans: Plasmodium falciparum, P. malariae, P. vivax
set up an environment conducive to parasite growth and and P. ovale. Most of the effort in developing a vaccine has
development. When zoites of T. gondii penetrate into host focused on P. falciparum, because it causes the most
cells, a tight junction is formed between the host cell and debilitating and lethal infections. Immunizing against
parasite at the site of penetration, inducing the release of malaria is complicated by the complexity of development
rhoptry contents. Dense granules are also released during in the human host, which involves several stages in red
and shortly after penetration. Once inside the cell, a space blood cells and in the liver: the sporozoite, the exoery-
develops between the host cell and the parasite; this is throcytic (EE) forms in the liver, and the asexual and sexual
called the parasitophorous vacuole. Multiplication of the erythrocytic forms. Initially, efforts to develop a human
parasite appears to be dependent upon proteins from malaria vaccine were directed against sporozoites or
rhoptries and dense granules becoming incorporated into erythrocytic merozoites. However, it was discovered that
the parasitophorous vacuole membrane and the develop- immunizing against sporozoites failed to protect against
ment of a tubulovesicular membrane network within the erythrocytic merozoite infection. If a single sporozoite
parasitophorous vacuole. escaped immune detection, then a clinical infection could
result.
It is now accepted that a multistage and multivalent
Immunity vaccine will be needed to achieve an efficacious malaria
vaccine. Compared to a single-stage antigen-based vac-
The development of immunity is dependent upon the cine, a multistage and multivalent vaccine would be more
outcome of a complex and dynamic interplay between the efficacious by inducing multiple layers of immunity.
host and the apicomplexan and is mediated through the Recent efforts have focused on developing genetically
host’s immune system. The immune system changes engineered vaccines consisting of genes encoding for as
adaptively in response to the activity and behaviour of many as nine stage-specific antigens of P. falciparum
the apicomplexan. corresponding to sporozoite, EE forms, and erythrocytic
Both antibody-dependent and cell-mediated (CMI) asexual and sexual stages. Preliminary studies have shown
mechanisms contribute to immunity in malaria as well as that such a multicomponent vaccine can induce immune
in other apicomplexans. Antigenic variation among responses that inhibit parasite development at multiple
malarial parasites results in considerable variation in their stages. Thus, an effective human malaria vaccine may be on
sensitivity to antibody-mediated inhibition. Both Ãã and the horizon.
T cells are important in malaria immunity. Although T Cryptosporidium parvum is a common source of diar-
cells may increase 40-fold or more in acutely infected rhoea in animals and usually causes a mild enteric disease
humans and also inhibit growth of parasites in vitro and in in normal humans, and leaves the host solidly immune to
vivo, their relative importance in protection or pathogeni- reinfection. On the other hand, severe, chronic infections
city is still not clear. Both CD4 1 and CD8 1 Ãã T cells may develop in immunocompromised hosts. Infection
respond to malarial infection, but the role of CD8 1 T cells induces specific IgM, IgG, IgA and IgE antibody
in blood stage infections appears to be limited. CD4 1 cells responses, and transient local and secretory IgA antibody
are of major importance and comprise at least two responses occur in the duodenum. Although the role of
functionally different subsets (TH1 and TH2), with TH1 specific antibodies remains unclear, decreases in oocyst
cells producing primarily interleukin 2 (IL-2) and inter- shedding in lambs and calves coincide with a rise in levels of
feron gamma (IFN-) and giving rise to protection early in coproantibodies, or antibodies in the intestinal contents.
infections. TH2 cells produce primarily IL-4 and are However, experimental studies in B-cell deficient mice do
primarily responsible for clearance of parasites late in the not suggest a protective role for either endogenous or
infection. In areas of high endemicity of malaria, most secretory antibodies. Also, high antibody titres of IgG and
humans have significantly elevated blood levels of IgE, but IgM were found in many AIDS patients with chronic C.
only approximately 5% of the IgE antibodies are directed parvum infections. Nevertheless, colostrum from cows
against P. falciparum, indicating that there is skewing of immunized against C. parvum decreases parasite burden
the underlying T-helper cell ratio in favour of TH2. IgE and is moderately efficacious against C. parvum when given
levels are highest in patients with cerebral malaria who also by mouth in AIDS patients with severe diarrhoea. Similar

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examples of transmission-blocking immunity have been Order Adeleina

studied in poultry coccidiosis and malaria in mankind and Macro- and microgamont usually associated in syzygy
have potential in the control of these diseases. during development; one to four microgametes; sporo-
Most primary infections with T. gondii in immunocom- zoites enclosed in an envelope, monoxenous or hetero-
petent hosts are clinically mild, but after the host recovers,
xenous. Adelea, Haemogregarina, Klossiella, Hepatozoon.
the parasite may remain encysted in the central nervous
system. Primary infection usually confers long-term Order Eimeriina
protection against reinfection and it is rare for seropositive
mothers to give birth to T. gondii-infected infants. Macro- and microgamonts develop separately; large
Humoral antibodies react against several surface as well number of microgametes formed; zygote nonmotile; no
as cytoplasmic antigens. However, the CMI response to T. syzygy; sporozoites typically enclosed in a sporocyst
gondii represents the major component of host immunity which, in turn, is enclosed in an oocyst; large number of
which can be divided into an innate and a parasite-specific microgametes; monoxenous or heteroxenous. Eimeria,
response. During acute infection, IL-2, produced by Isospora, Tyzzeria, Sarcocystis, Toxoplasma, Frenkelia,
macrophages accelerates the proliferation of natural killer Cyclospora.
(NK) cells that release IFN-a to stimulate the release of
nitrous oxide (NO). Although NO does not appear to be Order Cryptosporiina
involved in acute infections, it is important in controlling Macro- and microgamonts develop separately; 16 or fewer
parasite proliferation during chronic infections. After merozoites and microgametes; sporozoites enclosed in a
several days of infection, a parasite-specific T-cell response membrane that is either single or double; rhoptry-like
occurs, which provides long-term protection via the bodies in zoite, small conoid in some stages; no micro-
synergistic actions of CD4 1 and CD8 1 T cells. In nemes or polar ring; sporogony within the vertebrate host;
addition to IFN-g, a number of other cytokines are also retrofection an important part of the life cycle; endogenous
important mediators of the CMI response, in particular IL- forms usually about 5 mm and lying under the plasma
2, but IL-4, IL-6, IL-7 and IL-15 are also involved. membrane of enterocytes, but extracytoplasmically; para-
sites of birds, reptiles and mammals; monoxenous.
Cryptosporidium.
Major Taxa and Important Species
Class Haemosporea
Class Perkinsea Zoites have rhoptries, micronemes and polar ring, but lack
Order Perkinsida. One family, one genus, Perkinsus. a conoid; macro- and microgamonts develop separately,
Conoid forms an incomplete truncated cone; sexuality microgamont produces eight-flagellated microgametes;
absent; with flagellated zoospores that have an anterior zygote motile (ookinete); sporozoite naked; heteroxenous
vacuole; monoxenous. Perkinsus marinus. with merogony in vertebrates and sporogony in inverte-
brates; transmission by blood-sucking insects. Plasmo-
dium, Leucocytozoon, Haemoproteus.
Class Gregarinea
Conoid present in sporozoite for a period of time, Class Piroplasmea
converted to mucron upon entry into host cell; merogony
absent except in one group; gamonts large, extracellular, Zoites have rhoptries, micronemes, and polar ring, and
usually found in syzygy; gametes usually similar (isoga- some species have a conoid; parasites of erythrocytes and
mous); parasites of the digestive tract or body cavity of leucocytes where the organisms are small rod-like, piri-
invertebrates and some lower chordates; monoxenous. form, round or amoeboid forms; usually lacking micro-
Three orders: Archigregarinida, Eugregarinida, Neogre- tubules; locomotion by body flexion, gliding, or in sexual
garinida. Gregarina, Monocystis, Selenidium, Mattesia. stages by the large axopodium; sexual reproduction by
syngamy; heteroxenous with merogony in vertebrates and
gamogony and sporogony in invertebrates; sporozoites
Class Coccidea with single-membraned wall; ticks are the only known
All elements of the apical complex present; sporogony, vectors, but vectors of dactylosomatids are unknown.
merogony and gamogony present in all except two obscure Babesia, Theileria, Dactylosoma.
orders.
References
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Vector Biology, 2nd edn. San Diego: Academic Press.

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