Lizard Parasite Research Articles

Checklist of helminths from lizards and amphisbaenians (Reptilia, Squamata) of South America

A comprehensive and up to date summary of the literature on the helminth parasites of lizards and amphisbaenians from South America is herein presented. One-hundred eighteen lizard species from twelve countries were reported in the literature harboring a total of 155 helminth species, being none acanthocephalans, 15 cestodes, 20 trematodes and 111 nematodes. Of these, one record was from Chile and French Guiana, three from Colombia, three from Uruguay, eight from Bolivia, nine from Surinam, 13 from Paraguay, 12 from Venezuela, 27 from Ecuador, 17 from Argentina, 39 from Peru and 103 from Brazil. The present list provides host, geographical distribution (with the respective biome, when possible), site of infection and references from the parasites. A systematic parasite-host list is also provided.

Searching for virulence factors in the non-pathogenic parasite to humansLeishmania tarentolae

Leishmania protozoa are obligate intracellular parasites that reside in the phagolysosome of host macrophages and cause a large spectrum of pathologies to humans known as leishmaniases. The outcome of the disease is highly dependent on the parasite species and on its ascribed virulence factors and the immune status of the host. Characterization of the genome composition of non-pathogenic species could ultimately open new horizons in Leishmania developmental biology and also the disease monitoring. Here, we provide evidence that the lizard non-pathogenic to humans Leishmania tarentolae species expresses an Amastin-like gene, cysteine protease B (CPB), lipophosphoglycan LPG3 and the leishmanolysin GP63, genes well-known for their potential role in the parasite virulence. These genes were expressed at levels comparable to those in L. major and L. infantum both at the level of mRNA and protein. Alignment of the L. tarentolae proteins with their counterparts in the pathogenic species demonstrated that the degree of similarity varied from 59% and 60% for Amastin, 89% for LPG3 and 71% and 68% for CPB, in L. major and L. infantum, respectively. Interestingly, the A2 gene, expressed specifically by the L. donovani complex which promotes visceralization, was absent in L. tarentolae. These findings suggest that the lack of pathogenicity in L. tarentolae is not associated with known virulence genes such as LPG3, CPB, GP63 and Amastin, and that other factors either unique to L. tarentolae or missing from this species may be responsible for the non-pathogenic potential of this lizard parasite.

Parasitic infracommunities of the Aegean wall lizard Podarcis erhardii (Lacertidae, Sauria): isolation and impoverishment in small island populations

AbstractThe Aegean wall lizard Podarcis erhardii, is widely distributed across the islands of the Aegean Sea (Greece). While there exists a relatively substantial body of knowledge on the ecology and life history of the species, the parasite communities of the taxon remain almost completely unknown. Quantifying the composition of these communities in P. erhardii is not only important for autoecological reasons, but also because inter-island comparisons of this lizard's parasite communities can shed light on the factors that structure parasite diversity in general. Here we investigate the gastrointestinal parasite communities of P. erhardii populations occurring on 16 landbridge islands of the Sporades group in the NW Aegean Sea by examining the gastrointestinal tracts of 113 lizards. In all, 8 species of helminths were found: 1 Trematode (Paradistomum mutabile), 1 Cestode (Oochoristica sp.) and 6 Nematodes (Parapharyngodon micipsae, Parapharyngodon bulbosus, Parapharyngodon echinatus, Spauligodon sp., Abbreviata sp., and Skrjabinelazia sp.). The prevalence, mean intensity, and mean abundance of infection were respectively 63.71%; 6.01 (±11.71; range 1-90); and 3.57 (±9.5; range 0-90). Brillouin's index of diversity for the Sporades was 0.048 (±0.13; range 0-0.142). These values were lower than for most other mainland and insular lacertid populations, and suggest that the investigated island populations harbor very depauperate helminth communities. The severe impoverishment of the parasite communities and the differential persistence of generalist parasite species with simple life cycles is most likely the result of a combination of insular environmental conditions (spatial and temporal isolation, arid climate, small host population sizes) and host life history characteristics (diet, simple gastrointestinal tract architecture). The paucity of parasites in these relictual island populations suggests that small reptile populations fragmented by anthropogenic activities may not be able to sustain their native parasite communities over the long term.

Clonal diversity of a lizard malaria parasite, Plasmodium mexicanum, in its vertebrate host, the western fence lizard: role of variation in transmission intensity over time and space

Within the vertebrate host, infections of a malaria parasite (Plasmodium) could include a single genotype of cells (single-clone infections) or two to several genotypes (multiclone infections). Clonal diversity of infection plays an important role in the biology of the parasite, including its life history, virulence, and transmission. We determined the clonal diversity of Plasmodium mexicanum, a lizard malaria parasite at a study region in northern California, using variable microsatellite markers, the first such study for any malaria parasite of lizards or birds (the most common hosts for Plasmodium species). Multiclonal infections are common (50-88% of infections among samples), and measures of genetic diversity for the metapopulation (expected heterozygosity, number of alleles per locus, allele length variation, and effective population size) all indicated a substantial overall genetic diversity. Comparing years with high prevalence (1996-1998 = 25-32% lizards infected), and years with low prevalence (2001-2005 = 6-12%) found fewer alleles in samples taken from the low-prevalence years, but no reduction in overall diversity (H = 0.64-0.90 among loci). In most cases, rare alleles appeared to be lost as prevalence declined. For sites chronically experiencing low transmission intensity (prevalence approximately 1%), overall diversity was also high (H = 0.79-0.91), but there were fewer multiclonal infections. Theory predicts an apparent excess in expected heterozygosity follows a genetic bottleneck. Evidence for such a distortion in genetic diversity was observed after the drop in parasite prevalence under the infinite alleles mutation model but not for the stepwise mutation model. The results are similar to those reported for the human malaria parasite, Plasmodium falciparum, worldwide, and support the conclusion that malaria parasites maintain high genetic diversity in host populations despite the potential for loss in alleles during the transmission cycle or during periods/locations when transmission intensity is low.

Molecular identification of a Cryptosporidium saurophilum from corn snake (Elaphe guttata guttata)

Fecal samples from two snakes (Elaphe guttata guttata) have been examined by immunofluorescence test (IFT) and indicated Cryptosporidium spp. infection. Subsequent polymerase chain reaction (PCR), restriction fragment length polymorphism (RFLP) analysis and sequencing of small subunit (SSU) rRNA and actin genes enabled a distinct molecular characterization of the infecting organism as Cryptosporidium saurophilum. This is the first report about C. saurophilum infection in Elaphe guttata guttata. Cryptosporidium infections are common in reptiles and have been reported in at least 57 reptilian species (O’Donoghue 1995). Two main Cryptosporidium spp. are recognized in reptiles: Cryptosporidium serpentis is a gastric parasite mainly in snakes, and Cryptosporidium saurophilum is an intestinal parasite mainly in lizards (Morgan et al. 1999; Xiao et al. 2004a). Unlike other animals in which infection with Cryptosporidium spp. is usually self limiting in immunocompetent individuals, Cryptosporidium in reptiles is frequently chronic and sometimes lethal (Pasmans et al. 2007). C. serpentis infection in lizards is usually asymptomatic, whereas the infection in snakes frequently causes clinical disease (gastric hyperplasia, postprandial regurgitation and firm midbody swelling or chronic debilitating enteritis) and pathological changes (Brownstein et al. 1977; Kimbell et al. 1999; Xiao et al. 2004a; Cranfield et al. 1999). No pathological changes were found in the intestine and cloacae of adult lizards infected by C. saurophilum, but weight loss, abdominal swelling, and mortality occurred in some colonies of juvenile geckos (Eublepharis macularius) (Xiao et al. 2004b). Although C. saurophilum was originally described as a lizard parasite, it has been found in two captive snakes in Missouri (both snakes were heavily infected, and one had no clinical signs of diseases) and in three snakes (one Pituophis melanoleucuc sayi and two Pituophis ruthveni) in St. Louis Zoo (Xiao et al. 2004a, b). A group of six snakes (Elaphe obsolete, Chondrophyton viridis, Lampropeltis triangulum, Condoia asper, Pituophis melanoleucus) housed together with four lizards in the same room in Maryland also had C. saurophilum infections, but with much lower intensity than the infection of the four lizards, and all snakes were infected with multiple Cryptosporidium spp. (Xiao et al. 2004a). Studies on isolates recovered from wild and captive animals indicated that other and new Cryptosporidium spp. also exist in reptiles: a tortoise genotype identified in three tortoises, two new snake genotypes (one genotype identified in only one snake and the other genotype identified in six snakes), and another new Cryptosporidium genotype from a lizard, which was genetically distinct but was related to C. serpentis. Oocysts of the C. parvum bovine and mouse genotypes and C. muris found in some of the snakes and lizards were probably from rodents and mice ingested by these reptiles (Upton et al. 1989; Xiao et al. 2004a; Morgan et al. 1999). This possibility was supported by the fact that none of the animals with these oocysts had clinical signs and by the presence of organisms belonging to C. muris and the C. parvum mouse genotype in some of the feeder mice (Xiao et al. 2004a). In this study, we examined fecal samples from two snakes (Elaphe guttata guttata) and characterized the SSU rRNA and actin genes of Cryptosporidium-IFT positive samples by PCR-RFLP and DNA sequencing. Parasitol Res DOI 10.1007/s00436-007-0569-9

Clinical efficacy of antiparasite treatments against intestinal helminths and haematic protozoa in Gallotia caesaris (lizards)

Intestinal helminths and blood protozoa are common parasites of lizards. In captivity they can be a serious problem, but no information on treatments is available. In this study, several antiparasitic drugs were studied in Gallotia caesaris (Lacertida) from La Gomera, the Canary Islands, Spain. A lack of efficacy of chloroquine, and a decrease in haemogregarine (Apicomplexa) infection by atovaquone–proguanil was demonstrated. Furthermore, a positive effect of fenbendazole against intestinal nematode infection was observed. In the Canary Islands, Gallotia bravoana and Gallotia simonyi (Lacertida) are included in a captive breading recovery plan. The present results are useful in order to keep these animals in good condition and to control their parasites. Moreover, these results are important for reptiles kept as pets, due to the previous lack of knowledge of parasite management.

Identification of microsatellite markers in Plasmodium mexicanum, a lizard malaria parasite that infects nucleated erythrocytes

AbstractReptile and bird hosts of malaria parasites (Plasmodium) have nucleated erythrocytes. Infected blood thus contains a mix of abundant host and scant parasite DNA which has prevented identification of Plasmodium microsatellites. We developed a protocol for isolation of microsatellite markers for Plasmodium mexicanum, a parasite of lizards. The ATT repeat was common in the genome of P. mexicanum, but most (87%) of these repeats were exceptionally long (50–206 + repeats). Seven microsatellite markers with polymerase chain reaction primers are described. The protocol should allow discovery of microsatellites of malaria parasites (with AT‐rich genomes) infecting bird and reptile hosts.

Detection of a Malaria Parasite (Plasmodium mexicanum) in Ectoparasites (Mites and Ticks), and Possible Significance for Transmission

Two species of sandflies (Lutzomyia) are competent vectors of Plasmodium mexicanum, a malaria parasite of lizards. The very patchy distribution of sites with high P. mexicanum prevalence in the lizards, and often low or even nil sandfly density at such sites, provoked an evaluation of 2 common lizard ectoparasites, the tick Ixodes pacificus and the mite Geckobiella occidentalis, as potential passive vectors. Plasmodium sp.-specific polymerase chain primers were used to amplify a long segment of the mitochondrial cytochrome b gene that is unlikely to survive intact if the parasite cells are killed within a blood-feeding arthropod. The segment was strongly amplified from sandflies (the positive control for the method) from 1 to 96 hr postfeeding on an infected lizard. For ticks, the gene fragment was poorly amplified at 0 hr postfeed, and not amplified after 2 hr. In contrast, strong amplification of the parasite DNA was observed from mites from 0 to 20 hr postfeed, and weak amplification even at 96 hr.

Raillietiella morenoi sp. n. (Pentastomida) from Gallotia atlantica (Peters and Doria, 1882) (Lacertidae) in the Canary Islands

Raillietiella morenoi sp. n., a new cephalobaenid pentastomid found in the lungs of Gallotia atlantica collected in the Canary Islands, is described. The new species belongs to the sharp-tipped posterior-hook type. The annulus number, morphology, and dimensions of copulatory spicules and the dimensions of anterior and posterior hooks separate Raillietiella sp. n. from the other raillietiedid sharp-tipped posterior-hook species of small lizard parasites in Africa. The host character of endemic protected species of G. atlantica in Alegranza Island posed great difficulty in obtaining more parasite materials. However, more studies are required to state the variability of this species and its possible distribution in other species in the Canary Islands, as well as in other Gallotia spp.

Helminth Records from Eleven Species of Emoia (Sauria: Scincidae) from Oceania

As part of an ongoing study of the biogeography of helminth parasites of lizards from Oceania, 53 specimens of Emoia (11 species) were examined, as follows: E. atrocostata, E. boettgeri, E. caerulocauda, E. cyanogaster, E. cyanura, E. impar, E. nigra, E. nigromarginata, E. ponapea, E. sanfordi, E. trossula. One species of Digenea, Paradistomoides gregarium, and six species of Nematoda, Hedruris hanleyae, Maxvachonia chabaudi, Parapharyngodon maplestoni, Physalopteroides arnoensis, Spauligodon gehyrae, and Moaciria sp. indet., were found. These helminths have been reported previously from other lizard species. Seventeen new host records and eight new locality records are reported.

Rediscovery and redescription of Plasmodium pifanoi and description of two additional Plasmodium parasites of Venezuelan lizards.

Plasmodium pifanoi Scorza and Dagert B., known only from the type host, Ameiva ameiva, is redescribed from Kentropyx calcarata collected in Territorio Amazonas, Venezuela. Schizonts, 6.2 x 4.5 (4-8 x 3-6), produce on average 11.9 (7-16) merozoites. Gametocytes average 12.4 x 6.0 (8-16 x 4-10), with length x width (LW) 72.9 (52-112) and L/W 2.18 (1.1-3.3), and always contain 1-5 prominent vacuoles. Macrogametocytes in active infection are longer than microgametocytes, with greater LW, but gametocytes in chronic infection are not sexually dimorphic in dimension and are slightly smaller. Two additional malarial parasites are described from K. calcarata. Plasmodium lepidoptiformis has small schizonts, 4.6 x 3.2 (3-6 x 2.5-3), that produce 5.1 (4-8) merozoites and commonly resemble a butterfly in appearance. Gametocytes are elongate, 9.0 x 4.3 (7-10 x 3-6), with LW 38.3 (24-51) and L/W 2.2 (1.3-3.3), and sexually dimorphic, with macrogametocytes longer than microgametocytes, with greater LW. Plasmodium minasense calcaratae is characterized by very small, usually fan-shaped, schizonts. 3.4 x 2.6 (2.5-4.5 x 2.0-3.0), that produce 3.9 (3-4) merozoites. Gametocytes are spherical or ovoid, 6.7 x 5.0 (4.5-9.0 x 3.0-7.0), with LW 33.7 (15-54) and L/W 1.4 (1.0-2.3), with no sexual dimorphism in dimensions.

A molecular phylogeny of malarial parasites recovered from cytochrome b gene sequences.

A phylogeny of haemosporidian parasites (phylum Apicomplexa, family Plasmodiidae) was recovered using mitochondrial cytochrome b gene sequences from 52 species in 4 genera (Plasmodium, Hepatocystis, Haemoproteus, and Leucocytozoon), including parasite species infecting mammals, birds, and reptiles from over a wide geographic range. Leucocytozoon species emerged as an appropriate out-group for the other malarial parasites. Both parsimony and maximum-likelihood analyses produced similar phylogenetic trees. Life-history traits and parasite morphology, traditionally used as taxonomic characters, are largely phylogenetically uninformative. The Plasmodium and Hepatocystis species of mammalian hosts form 1 well-supported clade, and the Plasmodium and Haemoproteus species of birds and lizards form a second. Within this second clade, the relationships between taxa are more complex. Although jackknife support is weak, the Plasmodium of birds may form 1 clade and the Haemoproteus of birds another clade, but the parasites of lizards fall into several clusters, suggesting a more ancient and complex evolutionary history. The parasites currently placed within the genus Haemoproteus may not be monophyletic. Plasmodium falciparum of humans was not derived from an avian malarial ancestor and, except for its close sister species, P. reichenowi, is only distantly related to haemospordian parasites of all other mammals. Plasmodium is paraphyletic with respect to 2 other genera of malarial parasites, Haemoproteus and Hepatocystis. Explicit hypothesis testing supported these conclusions.

Non-pathogenic trypanosomatid protozoa as a platform for protein research and production

All currently existing eukaryotic protein expression systems are based on autonomous life forms. To exploit the potential practical benefits associated with parasitic organisms we have developed a new protein expression system based on Leishmania tarentolae (Trypanosomatidae), a protozoan parasite of lizards. To achieve strong transcription, the genes of interest were integrated into the small subunit ribosomal RNA gene. Expression levels obtained were up to 30 mg of recombinant protein per liter of suspension culture and increased linearly with the number of integrated gene copies. To assess the system's potential for production of post-translationally modified proteins, we have expressed human erythropoietin in L. tarentolae. The recombinant protein isolated from the culture supernatants was biologically active, natively processed at the N-terminus, and N-glycosylated. The N-glycosylation was exceptionally homogenous, with a mammalian-type biantennary oligosaccharide and the Man 3GlcNAc 2 core structure accounting for >90% of the glycans present. L. tarentolae is thus the first described biotechnologically useful unicellular eukaryotic organism producing biantennary fully galactosylated, core-α-1,6-fucosylated N-glycans.

First record of the mite Hirstiella diolii Baker (Prostigmata: Pterygosomatidae) from Australia, with a review of mites found on Australian lizards

AbstractWe report the first occurrence in Australia of an exotic mite parasite of lizards, Hirstiella diolii Baker (Prostigmata, Pterygosomatidae), and its association with iguanas at the Taronga Zoo in Sydney. We also report on the spread of the exotic snake mite Ophionyssus natricis (Gervais) to native populations of skinks, provide a key to the mites associated with Australian lizards and review how to distinguish chiggers from pterygosomatid mites.

Transmission success of the malaria parasite Plasmodium mexicanum into its vector: role of gametocyte density and sex ratio.

The life-cycle of Plasmodium depends on transmission of the parasite from the vertebrate host into its vector when the insect takes a bloodmeal. Transmission success may depend in part on the parasite's gametocyte density and sex ratio in the blood. P. mexicanum, a parasite of fence lizards in California, USA, exploits the sandfly Lutzomyia vexator as its vector. In experimental transmissions using naturally infected lizards as donors of blood, transmission success (measured as percentage of vectors infected and number of parasite oocysts on the insect's midgut) was positively related to gametocyte density, although density above 20/1000 erythrocytes did not improve transmission. Sex ratio (proportion of microgametocytes in the infection) was positively correlated with gametocyte density. Transmission improved with higher proportion of microgametocytes, but partial correlations revealed that this was a result only of higher gametocyte densities. These results agree with the theory of virulence and sex ratios because single clone infections should produce a more female-biased sex ratio and grow to the minimum parasitaemia that would maximize clonal transmission, whereas multiple clone infections will be closer to a 1:1 sex ratio and yield a higher parasitaemia when each clone competes for transmission to the vector.

Thubunaea dactyluris Sensu Fabio and Rolas, a synonym of Physalopteroides venancioi (Spirurida, Physalopteridae).

Specimens of Thubunaea dactyluris Karve, 1938 sensu Fabio and Rolas, 1974 from the lizard Ameiva ameiva of Brazil and types of Physalopteroides venancioi (Lent, Freitas, and Proença, 1946) from the toad Bufo paracnemis of Paraguay were re-examined. The male caudal papillae and the tooth arrangement demonstrated that these specimens were conspecific and are all assigned to P. venancioi. The occurrence of Physalopteroides dactyluris (Karve, 1938), a nematode parasite of lizards of India and Turkmenistan, is invalidated for Brazil.

Life history of a malaria parasite (Plasmodium mexicanum): independent traits and basis for variation.

Plasmodium mexicanum, a malaria parasite of lizards, exhibits substantial variation among infections in the life-history traits which define its blood-dwelling stages. Such variation in life histories among infections is common in Plasmodium and may influence the ecology and evolution of the parasite's transmission success and virulence. Insight into these issues requires identification of independent traits (some traits may be bound by developmental trade-offs) and the importance of genetic versus host effects producing the variation. We studied 11 life-history traits in 120 induced infections of P. mexicanum in its natural lizard host (20 each from six donor infections). The traits varied among infections and fell into three clusters: rate/peak (rate of increase and peak parasitaemia of asexuals and gametocytes), time (duration of pre-patent period and the infection's growth) and maturity (timing of first gametocytes). Thus, few life-history traits define an infection in the lizard's blood. Donor effects were significant for ten traits and two trait clusters (maturity was the exception) suggesting genetic differences among infections may influence the rate of increase and peak parasitaemia, but not the timing of the first production of gametocytes.

Leishmania (Sauroleishmania): A Comment on Classification

The classification of the Leishmania-like parasites of lizards has recently become the subject of sectional interests. In the absence of any objective criteria for classifying the group, two nomenclatures have come into current usage. Workers with interests in the natural history of the lizard parasites have emphasized the features that distinguish lizard and mammalian parasites and have tended to give the lizard parasites generic rank as the Sauroleishmania[1xSee all References][1]. Molecular biologists who have used the lizard parasites as a model organism have retained the lizard parasites within the genus Leishmania[2xPreviato, J.O. et al. Glycobiology. 1997; 7: 687–695Crossref | PubMed | Scopus (17)See all References][2].A number of studies, based on rRNA sequence comparisons, have been unable to resolve the phylogenetic relationship between the lizard parasite L. tarentolae and the mammalian Leishmania species[3xMarche, S. et al. Mol. Biochem. Parasitol. 1995; 71: 15–26Crossref | PubMed | Scopus (39)See all References][3]. However, two recent papers have provided the first DNA-based evidence that establishes the genetic relationships between the major groups of Leishmania species[4xNoyes, H.A. et al. J. Eukaryot. Microbiol. 1997; 44: 511–517Crossref | PubMed | Scopus (43)See all References, 5xCroan, D.G., Morrison, D.A., and Ellis, J.T. Mol. Biochem. Parasitol. 1997; 89: 149–152Crossref | PubMed | Scopus (121)See all References]. These studies present evidence from the RNA polymerase II, the DNA polymerase alpha and the small subunit rRNA genes and kinetoplast minicircles and they show that the lizard parasites diverged from the lineage leading to the L. (Leishmania) subgenus after this group separated from the New World parasites of the L. (Viannia) subgenus. This significant body of DNA evidence is inconsistent with a generic status for the lizard parasites but is consistent with acceptance of the subgenus L. (Sauroleishmania) as originally proposed by Saf'janova in 1982 (Ref. [1xSee all References][1]). Although the Leishmania-like parasites of the lizard gut were not included in these studies, there is already evidence to suggest that the lizard gut parasites are not close relatives of the lizard blood parasites and should not be included within the L. (Sauroleishmania), whatever taxonomic rank that group may have[1xSee all References][1].The correct classification of the lizard Leishmania is important. If these parasites are placed in a separate genus, there is a risk that they will be disregarded as an academic curiosity. If, as the available evidence suggests, they are derived from mammalian parasites, then the existing studies have real relevance to our understanding of the species that are human pathogens and there is a strong case for further work. The lizard parasites provide a useful model organism for studies of the molecular biology and biochemistry of the whole genus Leishmania, because they can be maintained in the laboratory with little risk to humans. Comparative studies of the lizard and mammalian parasites may shed further light on the mechanisms of the host–parasite relationship within the whole genus. Including the lizard parasites within the genus Leishmania should also prompt studies on the impact of these parasites on the epidemiology of the mammalian disease, since there is evidence of crossreactivity between lizard and mammalian parasite antigens in humans[1xSee all References][1].We urge parasitologists to adopt the subgenus L. (Sauroleishmania) Saf'janova, 1982, Ovezmukhammedov (1991)[1xSee all References][1] as the appropriate rank for the Leishmania from the blood of lizards and we look forward to the studies prompted by the renewed use of this designation.

Monophyletic origin of the genus Sauroleishmania

Summary A 1268-bp DNA fragment representing 25% of the RNA polymerase II largest subunit gene has been used to infer the phylogenetic relationships among four Sauroleishmania species, and their relatedness to the closely related genera Leishmania and Endotrypanum . Comparison of the nucleotide sequences showed that 94% of phylogenetically informative sites occurred at silent codon positions. Distance (neighbor-joining) and weighted parismony analysis of the nucleotide sequence alignment each produced alternate optimal trees which varied in the placement of L. (Viannia) braziliensis and the Sauroleishmania . The cost of branch swapping (number of steps in parsimony analysis) indicated that few steps were required to alternate between these topologies (representing 0.5% of total tree length). In contrast, distance and parsimony analysis of the predicted protein sequence yielded congruent results. The calculated homoplasy, retention and rescaled consistency indices of the protein sequence data indicated that homoplasy was significantly reduced in the protein sequence alignment as compared to the nucleotide sequence alignment. The protein-inferred phylogenies provide strong support for the monophyly of the Sauroleishmania and indicate that these parasites emerged subsequent to the separation of the Leishmania subgenera L. (Leishmania) and L. (Viannia) . The close affiliation between Leishmania and Sauroleishmania suggests that the lizard parasites might best be retained in the genus Leishmania .

Malarial Parasites of Lizards: Diversity and Ecology

Malarial Parasites of Lizards: Diversity and Ecology