Microsatellite mutations in litters of the Australian lizard Egernia stokesii

Seven tetranucleotide (AAAG) loci were analysed in a population (including 19 litters) of the Australian lizard Egernia stokesii. In an examination of 76 offspring we observed 13 mutations involving five loci. Two of our loci were highly mutable, with observed mutation rates of 2.7% and 4.2%, representing some of the highest mutations rates reported so far. A high proportion of mutations (46.2%) could not be assigned to changes involving only a single repeat, suggesting that mutations in at least two of the loci follow a multistep model. There was no significant bias of mutations leading to an increase or decrease in allele size; however, all multistep mutations involved a loss of repeats. These results add to increasing evidence casting doubt on microsatellite mutations being primarily single step mutations.     

Fine‐scale genetic structuring in a group‐living lizard,the gidgee skink (Egernia stokesii)

By constraining gene flow, group living and natal philopatry can result in fine‐scale genetic structure. Although the genetic structure of some group‐living lizards has been characterised, studies are few compared with those for group‐living bird and mammal species. The Egerniinae group of lizards exhibits a high diversity of social structures, making it a useful group for comparative studies of genetic structure across a broader range of social taxa. A well‐studied member of Egerniinae is Egernia stokesii, a lizard that forms long‐term pair bonds and stable social groups and exhibits natal philopatry and limited dispersal. Evidence exists for consistent E. stokesii social structure across seven close but disconnected rocky outcrops within a 40 × 10 km area. We used summary statistics, analysis of molecular variance, Bayesian clustering, and discriminant analysis of principal components to assess if E. stokesii exhibit a consistent pattern of fine‐scale genetic structure across the same seven outcrops. Due to E. stokesii social structure and constrained dispersal, we predicted significant genetic structuring – based on microsatellite markers – among outcrops. We found significant fine‐scale genetic structuring and evidence for two genetic clusters. We discuss features of E. stokesii biology and ecology that may explain our findings. Some rocky outcrops, and some social groups, contained lizards from both genetic clusters. An examination of the composition of mixed cluster social groups did not detect any notable patterns. Therefore, further work is necessary to identify how the observed patterns may have arisen. Future investigations in E. stokesii and other group‐living lizard species are likely to contribute greatly to our understanding of the genetic consequences of group living.     

High levels of genetic monogamy in the group-living Australian lizard Egernia stokesii

The Australian lizard Egernia stokesii lives in spatially and temporally stable groups of up to 17 individuals. We have recently shown that these groups are comprised of breeding partners, their offspring and, in some cases, highly related adults, providing the first genetic evidence of a family structure in any lizard species. Here we investigated the mating system of E. stokesii using data from up to eight polymorphic microsatellite loci and tested the hypothesis that breeding partners are monogamous both within and between mating seasons. Among 16 laboratory-born litters from field collected gravid females from two sites in South Australia, 75% had a single male parent and no male contributed to more than one litter, indicating a high level of genetic monogamy within a season. Additional analyses of field caught individuals, captured between 1994 and 1998, enabled assignment of parentage for 70 juveniles and subadults. These data showed that most young (88.6%) had both parents from within the same group and that high proportions of males (88.9%) and females (63.6%) have multiple cohorts of offspring only with the same partner. Our results suggest that monogamy both within and between seasons is a common mating strategy of E. stokesii and that breeding partners maintain stable associations together and with multiple cohorts of their offspring over periods of up to at least 5 years.     

Additional Fallisia spp. (Apicomplexa: Plasmodiidae) of Neotropical lizards

Fallisia biporcati n. sp. parasitises thrombocytes and lymphocytes of Anolis biporcatus and A. lionotus in Panama. Round or oval schizonts average 13.3 × 11.5 (10.5–16 × 9–13) m, with LW 153.8 (94–208) m², and produce 38.3 (28–60) merozoites. Gametocytes are variably shaped, from round or oval to nearly triangular or rectangular, and average 12.6 × 9.0 (10–15 × 6–12) m, with LW 113.1 (82–150) m² and L/W ratio 1.43 (1.0–2.2). Thrombocytes and lymphocytes of A. poecilopus in Panama are parasitised by F. poecilopi n. sp. Schizonts, oval to elongate in shape, 7.7 × 4.7 (5.5–9 × 3–6) m, with LW 36.5 (22–54) m², are filled with 31.0 (20–51) tiny nuclei or merozoites. Gametocytes are 10.1 × 8.0 (7.5–14 × 6–11) m, with LW 82.0 (45–132) m², round to elongate with L/W ratio 1.27 (1.0–1.6). F. thecadactyli n. sp. parasitises thrombocytes and lymphocytes of Thecadactylus rapicaudus in Panama and Venezuela. Oval, oblong, or triangular schizonts average 10.3 × 8.0 (7–13 × 5–12) m, with LW 86.6 (37–156) m², and produce 40.2 (26–61) merozoites. Gametocytes are round, oval, triangular or elongate, 10.4 × 7.0 (7–15 × 5–11) m, with LW 74.8 (40–154) m² and L/W ratio 1.51 (1.1–2.2). F. dominicensis n. sp. parasitises thrombocytes of A. cybotes on Hispaniola. Schizonts, 6.0 × 4.8 (4–8 × 3–7) m, with LW 29.1 (12–56) m², round, oval, elongate, oblong or lentiform in shape, produce 12.4 (8–22) merozoites. Gametocytes are 6.6 × 5.0 (5–9 × 4–7) m , with LW 33.8 (20–56) m², round, oval or elongate, and L/W ratio of 1.34 (1.1–2.0).     

Plasmodium spp. (Apicomplexa: Plasmodiidae) of the flying lizardDraco volans (Agamidae)

The agamid lizardDraco volans from Palawan, Republic of the Philippines, was found to be parasitised by threePlasmodium species:P. draconis n. sp.,P. volans n. sp. andP. vastator, Laird 1960. BothP. draconis andP. volans, but notP. vastator, were also present inD. volans from Sarawak. The species are readily distinguished by schizont size, merozoite number, and gametocyte size and shape.P. volans has schizonts approximately one-half the size of those ofP. draconis and produces 4–6 merozoites in comparison to 4–16 in the larger species. Gametocytes of both new species are predominantly oval in the Palawan sample and elongate in that from Sarawak.P. draconis gametocytes are approximately twice the size ofP. volans gametocytes, but on average no more than one-half the size of gametocytes ofP. vastator.     

Genetic evidence for a family structure in stable social aggregations of the Australian lizard Egernia stokesii

In this study we used data from six unlinked microsatellite loci to examine stable aggregations of Egernia stokesii, from a population in the southern Flinders Ranges of South Australia. We show that these aggregations are comprised of breeding partners, their offspring from two or more cohorts, and related adults, providing the first genetic evidence of a family structure in any lizard species. Despite this high level of relatedness within aggregations, most breeding pairs were unrelated and partners were less closely related to each other than they were to other potential within-group partners. Where individuals dispersed, both sexes usually moved to social groups close to their natal group. Although both sexes showed natal philopatry, there was some evidence that females in groups were more related than males in groups. These data suggest that an active choice of unrelated partners and male-biased dispersal may be the mechanisms used by E. stokesii to avoid inbreeding within groups.     

Thermoregulation in four Australian lizards of the genus Egernia (Sauria: Scincidae)

Mean thermal preferenda for the head and body were found to be similar in Egernia whitei, E. cunninghami, and E. bungana (range in T_H= 33.6^o -33.8^o C; range in T_B= 34.0^o-35.3^oC). However, those for E. majors were significantly lower (T_H= 31.1^o C, T_B=32.9^oC). Significant differences were found between mean head and body preferred temperatures in E. cunninghami and E. major. Higher temperatures were tolerated by the body than the head in all four species. Gaping was not observed to occur prior to cessation of respiration in any of the four species during radiant or solar heating. Based upon thermal preferenda and critical thermal maxima data, thermal safety margins for body temperature were calculated for E. whitei and E. cunninghami Postural orientation to the sun's rays increased and decreased heat gain. The seeking of shade was the most effective method employed by all four species for reducing head and body temperature.     

A Check-List of the Species of the Genus Leucocytozoon (Apicomplexa,Plasmodiidae)*

SYNOPSIS. A list is given of 67 named species of Leucocytozoon considered valid at present. Another list is given of 1 reptilian and 146 avian hosts of named species of Leucocytozoon. A 3rd list is given of 34 synonyms. Leucocytozoon has been named from about 0.7% of the approximately 8600 species of birds that exist. The following new taxonomic names are included: L. danilewskyi var. hirundinis Sergent & Sergent, 1905 is emended to L. hirundinis; Saurocytozoon tupinambi Lainson & Shaw, 1969 is changed to Leucocytozoon (Saurocytozoon) tupinambi (Lainson & Shaw, 1969).     

A Check-List of the Species of the Genus Haemoproteus (Apicomplexa,Plasmodiidae)*

SYNOPSIS. A list of the presumably valid named species of the genus Haemoproteus Kruse, 1890 (Apicomplexa, Plasmodiidae) is given. It includes 133 species, 5 varieties and 1 subspecies. One new species is named: H. serini n. sp. for the form in the canary Serinus canarius called H. zuenyoni by Sergent & Sergent (90). The following emendations are made: H. aegyptius for H. cellii var. aegyptius Mohammed, 1958; H. cellii for H. noctuae var. cellii Coatney & Roudabush, 1937; H. hirundinis for H. danilewskii var. hirundinis Sergent & Sergent, 1905; H. tinnunculus for H. danilewskyi var. tinnunculus Wasielewski & Wiilker, 1918; H. crumenius for H. crumenium (Hirst, 1905) Coatney, 1936; H. mathislegeri for H. herodiadis var. mathislegeri Mohammed, 1958; H. nebraskensis for H. noctuae var. nebraskensis Coatney & Roudabush, 1937; H. rileyi for Hoemoproteus rileyi Malkani, 1936; H. sauianae for H. sd-vianae Tendeiro, 1947; H. asturisdussumieri for H. asturis-dussumieri de Mello, 1935 emend. de Mello, 1937; H. xantholemae for H. xantholaemi Zargar, 1945. Of the 133 named species, 114 occur in birds, 16 in reptiles and 3 in amphibia; 14 orders and 47 families of birds are represented. The generic names Halteridium LabbC, 1894, Haemocystidium Castellani & Willey, 1904 and Simondia Garnham, 1966 are considered synonyms of Haemoproteus.     

Field studies on a social lizard: Home range and social organization in an Australian skink,Egernia major

Abstract Although most reptiles have polygynous mating systems without long‐term pair bonds, one lineage of large scincid lizards in Australia is exceptional in this respect. Reports of complex sociality in the genus Egernia led us to conduct the first radiotelemetric field study of a species within this group. Land mullets (Egernia major) are large (60 cm total length), viviparous lizards from rainforest habitats in south‐eastern Australia. To document the spatial ecology and social organization of this species, we captured 12 adult lizards in the Barrington Tops area of eastern New South Wales and implanted them with miniature radiotransmitters. The lizards were released at their sites of capture and located daily for the next 6 weeks. All of the radiotracked lizards had discrete home ranges of approximately 10 000 m², based around well‐defined core areas (approximately 2000–3000 m²). Females tended to move further, and to range over wider areas, than did males. All of the radiotracked lizards lived in social groups consisting of one or more adult males and females plus juveniles of all age classes. Subgroups were apparent within one group of five radio‐tagged lizards: individual animals consistently shared their shelter sites and home ranges with one or more specific individuals. Male/female pairings were more frequent than expected under the null hypothesis of random association among individuals. The data in the present study support anecdotal reports of pair bonds in E. major and support suggestions that the social systems of species in this genus are more complex than those of previously studied reptiles.     

Fallisia parasites (Haemosporidia: Plasmodiidae) from the flying lizard, Draco maculatus (Agamidae) in Thailand

Plasmodium (Fallisia) siamense n. sp. was described from the flying lizard Draco maculatus (Agamidae) collected near Bangkok, Thailand. Parasitic in thrombocytes, it produces 18-64 merozoites in schizonts that are larger than the usually oval gametocytes. Both gametocyte sexes showed a prominent nucleus, elongate in macrogametocytes and triangular in microgametocytes. This is the first Fallisia parasite described from a host outside the Neotropical Region, although presence of the genus in Australasia was reported 40 years ago, without formal taxonomic designation.     

The impact of nematode parasites on the behaviour of an Australian lizard,the gidgee skink <Emphasis Type="Italic">Egernia stokesii</Emphasis>

The Australian scincid lizard Egernia stokesii lives in social groups and is infected with two nematode species: Pharyngodon tiliquae and Thelandros trachysauri. This study asked whether those nematodes affected levels of lizard activity in field populations. In a laboratory colony, application of a combination of ivermectin and fenbendazole reduced nematode egg count in lizard scats after 12 weeks. In the field, the same doses of those antihelminthic drugs were applied to lizards in six social groups across three populations, and a saline control was given to lizards in six adjacent groups. Observations showed significant changes in behaviour between the two groups developing over 2 months. Drug-treated lizards spent more time basking and moved about for longer times during observation sessions. The results suggest that nematode infection altered host behaviour and reduced fitness. No influence of social group size was detected on the impact of parasitic nematodes.     

Two new species of Haemocystidium Castellani & Willey (Apicomplexa: Plasmodiidae) from Pakistani lizards, and the support their meronts provide for the validity of the genus

Haemocystidium papernai n. sp., a parasite of Agama nupta fusca in Sind, Pakistan produces large halteridial gametocytes in active infections that do not differ in morphology by sex. In chronic phase macrogametocytes are larger than microgametocytes. In Baluchistan, Agama nupta nupta is parasitised by Haemocystidium quettaensis n. sp. Gametocytes are smaller than those of H. papernai n. sp., round or oval in shape and heavily pigmented. Macrogametocytes are larger than microgametocytes and more heavily pigmented. Merogony of H. papernai n. sp. occurs in endothelium and connective tissue of lungs, heart, liver, spleen and femoral muscles. Meronts form as oval or rounded bodies that apparently are not preceded by the formation of pseudocytomeres that appear in the merogony of Haemoproteus spp. in snakes, turtles and birds. Meronts of Haemocystidium kopki were found in the lungs of the Pakistani gecko Teratoscincus scincus that were similar to those of H. papernai n. sp. The meronts of both species closely resemble phanerozoic meronts of saurian Plasmodium spp., and provide adequate justification for recognition of the genus Haemocystidium Castellani & Willey, 1904. Haemocystidium is distinguished from Plasmodium by the absence of an asexual cycle in circulating blood cells, and from Haemoproteus by meronts that do not form as pseudocytomeres. Sex ratios of H. papernai n. sp. varied from 22–47% microgametocytes, with no evident correlation with parasitemia, infection stage or month of collection, and H. kopki infections were comprised of 61–74% microgametocytes.     

艾美耳球虫对高原鼠兔繁殖的影响

寄生物对宿主繁殖的影响取决于宿主对当前繁殖值和剩余繁殖值的权衡。球虫为微型寄生物,而微型寄生物对宿主当前繁殖值的影响较剩余繁殖值要大。因此,本研究检验了寄生在高原鼠兔肠道内的艾美耳球虫可影响其当前繁殖的假设。在繁殖早、中、晚期,野外共观测高原鼠兔170只。结果表明,不同繁殖期感染率有显著差异。在繁殖中期,未感染雌性的妊娠率显著高于感染雌性。且未妊娠雌性较妊娠雌性有更高的感染强度,但在另外两个繁殖期没有发现此效应。在雄性中,任何繁殖期的感染强度和感染率与睾丸和附睾指数均无显著相关性,且感染和未感染球虫雄性睾丸及附睾指数无显著差异。此外,野外观测实验结果表明,感染雌性的胚胎重较未感染雌性显著降低,与野外感染对胚胎重量影响的实验结果相一致。说明艾美耳球虫感染可影响胚胎的发育。上述结果说明,艾美耳球虫对高原鼠兔繁殖的影响随繁殖期而有不同效应,且存在性别间差异,这种效应可能与不同性别间的繁殖对策有关。     

Coccidian parasites (Apicomplexa: Eimeriidae) of the endemic Florida snake Tantilla relicta Telford (Serpentes: Colubridae)

The endemic Florida snake Tantilla relicta Telford is parasitised by six species of coccidia. Caryospora tantillae n. sp. has nearly spherical oöcysts, 19.6 × 18.9 µm (16–22 × 16–21), with no polar body, and an oöcyst length/width ratio (shape index, SI) of 1.04 (1.00–1.11). Ovoidal sporocysts are 15.1 × 11.6 µm (12–17 × 10–13), with an SI of 1.30 (1.1–1.6), a single prominent Stieda body and a sporocyst residuum. Sporozoites are 14 × 2.5 µm. Caryospora relictae n. sp. also has nearly spherical oöcysts, 18.5 × 17.6 µm (16–22 × 15–20), but is distinguished by having a distinct polar body and a substieda body in sporocysts. The oöcyst SI is 1.05 (1.0–1.2). Sporocysts are ovoidal, 14.8 × 11.4 µm (12–17 × 10–16), with an SI of 1.29 (1.1–1.5) and a sporocyst residuum. Sporozoites are 10–14 × 2–2.5 µm. Isospora cf. wilsoni has ovoidal to spherical oöcysts without a micropyle, measure 17.4 × 17.1 µm (15–19 × 15–19), and have an SI of 1.02 (1.00–1.07). Oöcysts have no polar granule or oöcyst residuum. Sporocysts are ovoidal, 11.4 × 8.2 µm (9–14 × 7–10), with an SI of 1.39 (1.18–1.62). Both a Stieda body and sporocyst residuum are present, but a substieda body, characteristic of I. wilsoni as described, was not observed. Sporozoites are 7–8 × 2.5–3 µm. An Eimeria sp., of which the sample was too small to justify taxonomic designation, has nearly spherical oöcysts without micropyle, a size of 22.5 × 20.9 µm (22–23 × 20–22), an SI of 1.08 (1.00–1.12) and no polar granule. Sporocysts measure 12.5 × 8.9 µm (11–14 × 7–11) and have an SI of 1.44 (1.18–1.87), a sporocyst residuum and no Stieda body. Sporozoites are 13 × 5 µm. Eimeria amphorae n. sp. has ovoidal to spherical oöcysts without a micropyle, a size of 24.8 × 22.3 µm (23–27 × 20–26), an SI of 1.12 (1.02–1.25) and no polar granule or oöcyst residuum. Sporocysts are elongate, 18.4 X 8.0 µm (17-22 × 6-9.5), with an SI 2.33 (1.79–3.00) and no discernible Stieda body or sporocyst residuum. Sporocyst shape differs by orientation, rectangular to amphora-like, tapering abruptly to a blunt point; sporozoites large, 15.5 × 4.2 µm. A biliary parasite, Eimeria pleistocenensis n. sp., has ovoidal to elongate oöcysts, without micropyle, with a size of 27.8 × 19.4 µm (23–32 × 14–23), with an SI of 1.44 (1.17–1.82) and an oöcyst residuum but no polar granule. Sporocysts ovoidal, 10.7 × 8.6 (9–13 × 8–10), with an SI of 1.24 (1.05–1.37), a sporocyst residuum, but without Stieda body; sporozoites 8.8 × 2.7 µm. Similar oöcysts from other Tantilla relicta populations, T. relicta neilli in north Florida T. relicta pamlica in southeastern Florida and T. relicta relicta in south-central Florida, have dimensions that lie within the range of variation in the type population and identically shaped sporocysts that suggest conspecificity with E. pleistocenensis, and presence of the species in the precursor to Tantilla relicta prior to the second interglacial period.     

Redescription of Haemoproteus mesnili (Apicomplexa: Plasmodiidae) and its meronts, with description of a second haemosporidian parasite of African cobras

Haemoproteus mesnili (Bouet 1909) Wenyon 1926 is redescribed from the spitting cobra, Naja nigricollis nigricollis, of Tanzania. Mature gametocytes in the acute phase of infection averaged 17.7 X 7.3 jim, with LW 128.1 jim-, and L:W ratio 2.52. Nuclei were visible in both sexes. Both sexes were heavily pigmented, with 31-62 black granules dispersed in macrogametocytes; 20-46 granules were often clumped or concentrated near ends of microgametocytes. The halteridial form was present in 28% of active-phase gametocytes, but in only 8% of those in chronic phase. A few large, possibly first generation, meronts were present in cardiac muscle; uninucleate parasites within parasitophorous vacuoles in splenic cells produced small rounded or ovoid meronts, 12.2 x 9.6 microm, with 12-16 deeply basophilic, square-to-rectangular cytomeres. Meronts with 17-32 cytomeres were 16.9 x 11.9 microm. Meronts, 20 x 16 to 26 x 22 microm, contained 51-57 cytomeres. Mature meronts were ovoid, 13.7 x 11.5 microm, with many rounded merozoites. Haemoproteus balli n. sp, found in an Egyptian cobra, Naja haje haje of Kenya, differs from H. mesnili in average gametocyte dimensions, 10.8 x 7.7 microm; LW, 83.2 microm2; L/W ratio, 1.42; absence of halteridial forms; sparse pigmentation (3-10 granules); and presence of a broad peripheral band, apparently chromatin, along one side of microgametocytes.     

The impact of habitat fragmentation on dispersal of Cunningham's skink (Egernia cunninghami): evidence from allelic and genotypic analyses of microsatellites

The effects of habitat fragmentation on processes within and among populations are important for conservation management. Despite a broad spectrum of lifestyles and the conservation significance of many reptiles, very little work on fine-scale population genetics has been carried out on this group. This study examines the dispersal patterns of a rock crevice-dwelling lizard, Cunningham's skink (Egernia cunninghami), in a naturally vegetated reserve and an adjacent deforested site. Both genotypic and genic approaches were employed, using microsatellite loci. The spatial organization of individuals with respect to pairwise relatedness coefficients and allele frequencies, along with assignment tests, were used to infer dispersal characteristics for both sexes in a natural and a cleared area. The distribution of relatedness in both habitats was spatially structured, with E. cunninghami showing high pairwise relatedness within their rocky retreat sites. Analysis of relatedness over different spatial scales, spatial autocorrelation of alleles and assignment tests, all indicated that both sexes in the cleared area show less dispersal than their counterparts in the reserve. Furthermore, deforestation may inhibit female dispersal to a greater extent than that of males. The geographical structuring of allele frequencies for adults in the cleared area, but not the reserve, indicates that habitat fragmentation has the potential to alter at least the microevolution of E. cunninghami populations.