Population synchrony within and among Lepidoptera species in relation to weather, phylogeny, and larval phenology

Abstract.  1. The population dynamics of native herbivore species in central Appalachian deciduous forests were studied by analysing patterns of synchrony among intra- and interspecific populations and weather.
2. Spatial synchrony of 10 Lepidoptera species and three weather variables (minimum temperature, maximum temperature, precipitation) from 12 sites was measured using cross-correlation functions relating levels of synchrony to the distance separating each set of populations. Based on both the pattern of synchrony and the region-wide cross-correlation coefficients, Lepidoptera species appear to be synchronised, at least in part, by local weather conditions.
3. Interspecific cross-correlations were calculated for all sympatric species pairs and trends in interspecific synchrony were related to phylogenetic relatedness, life-history timing, and weather. Interspecific synchrony was highest among species whose larvae were present during the same time of the season, but there was no relationship between interspecific synchrony and phylogenetic affinity.
4. Spatial synchrony of weather variables was significantly related to both species of some interspecific pairs, indicating weather as a potential mechanism involved in synchronising populations of different species.     

PERIODIC THERMAL STRESS: OPTIMAL SPACING PREDICTED BY A SIMPLE SET-BACK MODEL OF CELL DIVISION SYNCHRONIZATION

Digital computer simulations have been used to make quantitative predictions based on a simple set-back model of cell division synchronization. According to the model appropriate thermal stress reverses progress within a segment of the division cycle called the set-back interval. In the simulations normally distributed cell-to-cell variations in division cycling rate between periodic thermal shocks were produced with the Monte Carlo method.
The simulations have shown that reasonably good synchronization with the single shock per division strategy requires a relatively long set-back interval and small cell-to-cell variations in rate of progress through the division cycle. The simulations have shown that the degree of synchrony produced by such periodic shocks is highly dependent on the time interval between shocks—with a series of as many as seven shocks inappropriately spaced producing less synchrony than a single shock! The optimal time interval between successive thermal shocks was found approximately equal to the mode division cycle time at synchrony equilibrium multiplied by 1 plus half the fraction of the division cycle occupied by the set-back interval. Position of the set-back interval within the division cycle had little effect on synchrony at the end of the final shock.     

Spatial variability in prey phenology determines predator movement patterns and prey survival

Species have phenological variation among local habitats that are located at relatively small spatial scales. However, less studies have tested how this spatial variability in phenology can mediate intra-/inter-specific interactions. When predators track phenological variation of prey among local habitats, survival of prey within a local habitat strongly influenced by phenological synchrony with their conspecifics in adjacent habitats. Theory predicts that phenological synchrony among local habitats increases prey survival in local habitat within spatially structured environments because the predators have to make a habitat choice for foraging. Consequently, total survival of prey at regional scale should be higher. By using a spatially explicit field experiment, we tested above hypothesis using a prey–predator interaction between tadpole (Rhacophorus arboreus) and newt (Cynops pyrrhogaster). We established enclosures (≈regional scale) consisting of two tanks (≈local habitat scale) with different degree of prey phenological synchrony. We found that phenological synchrony of prey between tanks within each enclosure decreased the mean residence time of the predator in each tank, which resulted in higher survival of prey at a local habitat scale, supporting the theoretical prediction. Furthermore, individual-level variation in predator residence time explained the between-tank variation in prey survival in enclosures with phenological synchrony, implying that movement patterns of the predator can mediate variation in local population dynamics of their prey. However, total survival at each enclosure was not higher under phenological synchrony. These results suggest the importance of relative timing of prey phenology, not absolute timing, among local habitats in determining prey–predator interactions.     

Within-pair testosterone covariation and reproductive output in Greylag Geese Anser anser

In monogamous animals, reproductive success may vary considerably among pairs. To study this variation, we focussed on reproductive events and the circannual hormonal co-ordination within pairs during the reproductive cycle. Testosterone was chosen as covariable for both sexes because of its mediator function between behaviour and physiology. In a flock of free-living Greylag Geese Anser anser , individual faecal samples were collected weekly from 23 pairs over a complete annual cycle. From the faeces, equivalents of testosterone, oestrogen and other steroid hormones were analysed by enzyme immunoassay. In contrast to correlations between male and female testosterone, no correlations were found between oestrogen-oestrogen or oestrogen-testosterone. Therefore, only testosterone (T) is considered here. Sex-specific differences in T were in amplitude rather than in the annual timing of maxima and minima. However, the annual timing varied to some degree between individuals within sexes. Therefore, we examined the degree of annual testosterone correlation (TC) within pairs. Pairs that nested had significantly higher TC over the year than pairs that did not. The higher the within-pair TC, the larger the clutches and the heavier the eggs. Also, TC in the year investigated was positively and significantly correlated with the pairs' long-term reproductive output. No correlation was found between TC and the duration of the pair-bond, individual age, or age difference from pair partner. We suggest that TC is a measure of behavioural synchrony and, therefore, pair-bond quality. We consider whether within-pair TC results from mate choice.     

Leaf synchrony and insect herbivory among tropical tree habitat specialists

Growth defense tradeoff theory predicts that plants in low-resource habitats invest more energy in defense mechanisms against natural enemies than growth, whereas plants in high-resource habitats can afford higher leaf loss rates. A less-studied defense against herbivores involves the synchrony of leaf production, which can be an effective defense strategy if leaf biomass production exceeds the capacity of consumption by insects. The aim of this study was to determine whether leaf synchrony varied across habitats with different available resources and whether insects were able to track young leaf production among tree habitat specialists in a tropical forest of French Guiana. We predicted that high-resource habitats would exhibit more synchrony in leaf production due to the low cost and investment to replace leaf tissue. We also expected closer patterns of leaf synchrony and herbivory within related species, assuming that they shared herbivores. We simultaneously monitored leaf production and herbivory rates of five pairs of tree species, each composed of a specialist of terra firme or white-sand forests within the same lineage. Our prediction was not supported by the strong interaction of habitat and lineage for leaf synchrony within individuals of the same species; although habitat specialists differed in leaf synchrony within four of five lineages, the direction of the effect was variable. All species showed short time lags for the correlation between leaf production and herbivory, suggesting that insects are tightly tracking leaf production, especially for the most synchronous species. Leaf synchrony may provide an important escape defense against herbivores, and its expression appears to be constrained by both evolutionary history and environmental factors.     

Maintenance of an Intact Human Immunodeficiency Virus Type 1 vpr Gene following Mother-to-Infant Transmission

The vpr sequences from six human immunodeficiency virus type 1 (HIV-1)-infected mother-infant pairs following perinatal transmission were analyzed. We found that 153 of the 166 clones analyzed from uncultured peripheral blood mononuclear cell DNA samples showed a 92.17% frequency of intact vpr open reading frames. There was a low degree of heterogeneity of vpr genes within mothers, within infants, and between epidemiologically linked mother-infant pairs. The distances between vpr sequences were greater in epidemiologically unlinked individuals than in epidemiologically linked mother-infant pairs. Moreover, the infants’ sequences displayed patterns similar to those seen in their mothers. The functional domains essential for Vpr activity, including virion incorporation, nuclear import, and cell cycle arrest and differentiation were highly conserved in most of the sequences. Phylogenetic analyses of 166 mother-infant pairs and 195 other available vpr sequences from HIV databases formed distinct clusters for each mother-infant pair and for other vpr sequences and grouped the six mother-infant pairs’ sequences with subtype B sequences. A high degree of conservation of intact and functional vpr supports the notion that vpr plays an important role in HIV-1 infection and replication in mother-infant isolates that are involved in perinatal transmission.     

Population synchrony of a native fish across three Laurentian Great Lakes: evaluating the effects of dispersal and climate

Climate and dispersal are the two most commonly cited mechanisms to explain spatial synchrony among time series of animal populations, and climate is typically most important for fishes. Using data from 1978–2006, we quantified the spatial synchrony in recruitment and population catch-per-unit-effort (CPUE) for bloater (Coregonus hoyi) populations across lakes Superior, Michigan, and Huron. In this natural field experiment, climate was highly synchronous across lakes but the likelihood of dispersal between lakes differed. When data from all lakes were pooled, modified correlograms revealed spatial synchrony to occur up to 800 km for long-term (data not detrended) trends and up to 600 km for short-term (data detrended by the annual rate of change) trends. This large spatial synchrony more than doubles the scale previously observed in freshwater fish populations, and exceeds the scale found in most marine or estuarine populations. When analyzing the data separately for within- and between-lake pairs, spatial synchrony was always observed within lakes, up to 400 or 600 km. Conversely, between-lake synchrony did not occur among short-term trends, and for long-term trends, the scale of synchrony was highly variable. For recruit CPUE, synchrony occurred up to 600 km between both lakes Michigan and Huron (where dispersal was most likely) and lakes Michigan and Superior (where dispersal was least likely), but failed to occur between lakes Huron and Superior (where dispersal likelihood was intermediate). When considering the scale of putative bloater dispersal and genetic information from previous studies, we concluded that dispersal was likely underlying within-lake synchrony but climate was more likely underlying between-lake synchrony. The broad scale of synchrony in Great Lakes bloater populations increases their probability of extirpation, a timely message for fishery managers given current low levels of bloater abundance.     

Common growth signal and spatial synchrony of the chronologies of tree-rings from pines in the Baltic Sea region over the last nine centuries

The common growth signal and spatial synchrony of nine chronologies of tree rings (Pinus sylvestris L.) from the Baltic Sea region were studied using moving correlations with time windows of 50 and 100 years. Drifts from synchrony to asynchrony and again back to synchrony across several centuries were observed. The chronologies showed higher (or lower) coefficients of correlations and correspondingly gentler (or steeper) declines in spatial synchrony in certain periods. In accordance with research into other ecological phenomena, the origin of the spatial synchrony was found to relate to the intensity of the westerly circulation over the Atlantic and Europe as described by the North Atlantic Oscillation (NAO) index. The tree rings from all of the analysed subregions were found to be wider, identifying faster periods of growth, when there was a positive NAO during the extended winter season from December to March that preceded the period of growth. Previous studies showed that the NAO index can be linked with the growth of tree rings because of factors in the local climate that affect growth, such as temperature and precipitation. Moreover, the spatial synchrony showed multidecadal fluctuations that were correlated with variations of a similar scale in the reconstructed NAO index. However, a high degree of spatial synchrony was observed during the 20^th century in the presence of values of NAO index that were strongly negative. A more detailed view was provided by maps of the correlations in which spatial changes were demonstrated in the growth of pines between the 12^th and the 20^th century. These results could be used as background information in the development of improved proxy-based reconstructions of the NAO index (or other similar meteorological variables), which could also be inferred from tree ring data relating to earlier centuries.     

Relationship between the phases of mitosis in the cells of chimeric loach embryos

The loach embryos differing in age by 0.5 tau0 (tau0--duration of the mitotic cycle during the synchronous cleavage divisions) were combined in pairs at the stage of 4--8 blastomeres to elucidate the role of intercellular relationships in the preservation of cell division synchrony during cleavage. Following the incubation during 3 tau0 the mean value of interval between mitotic phases in the pairs of combined embryos somewhat decreased but this phenomenon cannot be considered as the proof of cell division synchronization due to the intercellular relationships since it was observed to the same extent in all groups of chimaeric embryos irrespective of the degree of fusion as well as in the control embryos.     

棘尾虫无小核镜像骈体的获得及其生殖行为的实验观察

用在分裂期进行显微手术的方法和在陈旧培养中诱导的方法,从贻贝棘尾虫天然无小核系S_(10)中获得了无小核镜像骈体。对骈体的各种生殖行为进行了观察,并用孚尔根和改进的黑色素染色技术揭示了它们的细胞学细节。发现在本文描述的无小核骈体和过去报告的有小核骈体之间,在二分裂和生理再生上没有显著区别。无小核骈体和正常单体S_7之间的接合命运则是多样化的。在630个这样的接合对中,有257对发生了真正的接合并形成接合后体;有153对接合不久包囊化;在其余的220对中,一个接合体吸收了另一个,变成营养型的单体和有小核的骈体。前者占总吸收者的96.8％,后者只占3.2％。对接合和吸收两种情况的细胞学事件进行了详细观察。无小核骈体中的两组细胞质和双套大核能吸引其配偶中的小核,形成自己新的双套大核胚基及小核胚基,在本文的讨论中受到充分注意。     

Synchrony of Nuclear Replication in Individual Hyphae of Aspergillus nidulans

The synchrony of nuclear replication in individual, multinucleate hyphae of Aspergillus nidulans has been investigated. Samples were taken from cultures of germinating conidiospores, and the relative frequency of hyphae containing two to eight nuclei was determined. Because the conidiospores are mononucleate, complete synchrony will yield populations of hyphae containing only 2ⁿ nuclei, n being the number of doublings after germination. The appearance of hyphae with total numbers of nuclei other than 2ⁿ will indicate lack of synchrony. The relative frequency of hyphae not having 2ⁿ nuclei will depend on the degree of synchrony in the individual hyphae; numerical aspects of this relation are discussed. In two different strains, replication of the nuclei in any one hypha was highly synchronized when the dry weight doubling time was 1.4 to 1.8 hr. As the doubling time was made longer by changing the nitrogen or carbon source, synchrony was progressively lost. At the slowest growth rate tested, the interval between the division of the fastest and the slowest nucleus equaled 48% of the dry weight doubling time. The active replication of some nuclei in a hypha where other nuclei were resting suggested that nuclear duplication in this eukaryotic organism may be controlled by specific initiators.     

The Zinc Finger Protein Zfr1p Is Localized Specifically to Conjugation Junction and Required for Sexual Development in Tetrahymena thermophila

Conjugation in Tetrahymena thermophila involves a developmental program consisting of three prezygotic nuclear divisions, pronuclear exchange and fusion, and postzygotic and exconjugant stages. The conjugation junction structure appears during the initiation of conjugation development, and disappears during the exconjugant stage. Many structural and functional proteins are involved in the establishment and maintenance of the junction structure in T. thermophila. In the present study, a zinc finger protein-encoding gene ZFR1 was found to be expressed specifically during conjugation and to localize specifically to the conjugation junction region. Truncated Zfr1p localized at the plasma membrane in ordered arrays and decorated Golgi apparatus located adjacent to basal body. The N-terminal zinc finger and C-terminal hydrophobic domains of Zfr1p were found to be required for its specific conjugation junction localization. Conjugation development of ZFR1 somatic knockout cells was aborted at the pronuclear exchange and fusion conjugation stages. Furthermore, Zfr1p was found to be important for conjugation junction stability during the prezygotic nuclear division stage. Taken together, our data reveal that Zfr1p is required for the stability and integrity of the conjugation junction structure and essential for the sexual life cycle of the Tetrahymena cell.     

Scale-dependent spatial population dynamics of gall-makers on oak

The patterns of synchrony in the population fluctuations of six species of gall‐makers on oak (Hymenoptera, Cynipidae and Diptera, Cecidomyiidae) were analyzed over a small‐scale transect (8 km) and a large‐scale transect (500 km). Gall‐maker species differed in their degree of synchrony. At the small scale some species showed synchrony among local sites, whereas at the large scale, with one exception, population fluctuations of all species were largely independent. The patterns of synchrony differed between the two spatial scales. At the small scale a considerable degree of synchrony was found among sites for two species, Cynips divisa and Neuroterus quercusbaccarum, whereas at the large scale no synchrony was seen for these species. For one species, Macrodiplosis volvens, the fluctuations were asynchronous at both the small and large scales. At the large scale, synchrony among sites was found for one species: the fluctuations of Neuroterus anthracinus were largely synchronous at both scales (i.e. over several hundred kilometers). Distance‐dependent synchronies (i.e. decreasing synchrony with increasing distance) were found for only one species, Neuroterus anthracinus. In summary, the levels of synchrony in the population fluctuations of these insects differed among species and were scale‐dependent. Scaling up from the small scale to the large scale does not seem appropriate.     

The time and duration of the premeiotic inteprhase in microsporocytes ofTrillium kamtschaticum

The time and duration of each phase of the premeiotic interphase were determined in microsporocytes of two clones (S and K clones) ofTrillium kamtschaticum. After collectionTrillium plants were stored at 3 C or 7 C prior to completion of premeiotic mitosis in archesporial cells. For autoradiography, cells were explanted in the presence of³H-thymidine to identify the interval of the premeiotic DNA synthesis. Approximate durations of the G₁, S and G₂ phases for the K clone stored at 3 C were estimated to be 12, 12 and 14 days, respectively. The interval of premeiotic development was markedly different between clones. A high degree of synchrony in meiotic development, which is usually observed within anthers up to late meiotic prophase, was confirmed at the S phase, suggesting that synchrony is established during the G₁ interval.     

Environment,but not migration rate,influences extinction risk in experimental metapopulations

Ecological theory suggests that several demographic factors influence metapopulation extinction risk, including synchrony in population size between subpopulations, metapopulation size and the magnitude of fluctuations in population size. Theoretically, each of these is influenced by the rate of migration between subpopulations. Here we report on an experiment where we manipulated migration rate within metapopulations of the freshwater zooplankton Daphnia magna to examine how migration influenced each of these demographic variables, and subsequent effects on metapopulation extinction. In addition, our experimental procedures introduced unplanned but controlled differences between metapopulations in light intensity, enabling us to examine the relative influences of environmental and demographic factors. We found that increasing migration rate increased subpopulation synchrony. We failed to detect effects of migration on population size and fluctuations in population size at the metapopulation or subpopulation level, however. In contrast, light intensity did not influence synchrony, but was positively correlated with population size and negatively correlated with population fluctuation. Finally, synchrony did not influence time to extinction, while population size and the magnitude of fluctuations did. We conclude that environmental factors had a greater influence on extinction risk than demographic factors, and that metapopulation size and fluctuation were more important to extinction risk than metapopulation synchrony.     

Time-matched grooming in female primates? New analyses from two species

The parcelling model of reciprocity predicts that grooming partners will alternate between giving and receiving grooming within grooming bouts, and that each partner will perform approximately as much grooming as it receives within each bout (‘time matching’). Models of allogrooming based on biological markets theory predict that individuals of lower dominance rank will exchange grooming for tolerance from high-rankers, and therefore an inverse relation will be found between grooming partners' dominance rank distance and how closely they match each other's grooming contributions within each bout. We used weighted logistic regression and weighted least-squares regression to test these predictions using data from female white-faced capuchins, Cebus capucinus, and bonnet macaques, Macaca radiata. Only 5-7% of macaque grooming bouts, and 12-27% of capuchin grooming bouts, were reciprocated. However, (1) the duration of grooming by the first groomer significantly predicted whether the groomee would reciprocate at all, and (2) when bouts were reciprocated, the duration of grooming by the first groomer significantly predicted the duration of grooming by the second groomer. Grooming was most balanced among females of similar dominance ranks. Both the time-matching and rank-related effects were weak, although significant. These results indicate that although some form of time matching may be a general characteristic of grooming in female-bonded primate species, time matching accounts for relatively little of the variation in the distribution of grooming within bouts. We also draw attention to weighted regression as a technique that avoids pseudoreplication while using all available data.     

Analysis of the schedule of DNA replication in heat-synchronized Tetrahymena

The temporal schedule of DNA replication in heat-synchronized Tetrahymena was studied by autoradiographic and cytofluorometric methods. It was shown that some cells, which were synchronized by selection of individual dividing cells or by temporary thymidine starvation, incorporated [³H]thymidine into macronuclei in a periodic fashion during the heat-shock treatment. It was concluded that supernumerary S periods occurred while cell division was blocked by high temperature. The proportion of cells which initiated supernumerary S periods was found to be dependent on the duration of the heat-shock treatment and on the cell cycle stage when the first heat shock was applied. Cytofluorometric measurements of Feulgen-stained macronuclei during the heat-shock treatment indicated that the DNA complement of these cells was substantially increased and probably duplicated during the course of each S period. Estimates of DNA content also suggested that the rate of DNA synthesis progressively declined during long heat-shock treatments. These results indicate that the mechanism which brings about heat-induced division synchrony is not an interruption of the process of DNA replication. Further experiments were concerned with the regulation of DNA synthesis during the first synchronized division cycle. It was shown that participation in DNA synthesis at this time increased as more cells were able to conclude the terminal S period during the preceding heat-shock treatment. It is suggested that a discrete period of time is necessary after the completion of DNA synthesis before another round of DNA synthesis can be initiated.     

The influence of age on the use of potential enrichment objects and synchronisation of behaviour of pigs

This experiment aimed to identify how a pig's age affected the extent and synchrony of use of different environmental enrichment materials, and how this use changed over time. Ten diverse novel objects were each presented to three replicate litters of 3 weeks of age (sucklers) and three replicate groups of three animals of 5 (weaners) weeks and 13 (growers) weeks of age. Video recordings were made of the pigs’ behaviour over a period of 5 days and subsequently analysed for activity, inactivity and object directed behaviour of three animals per group on days 1 and 5. The observed performance of any given behaviour, when at least one other member of the group was also performing that behaviour, was compared with the probability that such concurrence occurred by chance and these results were used to calculate the degree of synchronisation. Gender had no effect on the duration of object use or approach latency. Growers displayed a shorter latency to approach the objects initially compared to sucklers and weaners (mean seconds; 2153 versus 2660 versus 980 for sucklers, weaners and growers, respectively, S.E.D. 542, p = 0.007). Sucklers used the objects to a much lesser extent than either the weaners or growers (mean duration object interaction (s); 129 versus 1253 versus 1412 for sucklers, weaners and growers, respectively, S.E.D. 93, p < 0.001). Overall object use decreased between days 1 and 5 (mean duration object interaction (s); 1326 versus 536 for days 1 and 5, respectively, S.E.D. 76, p < 0.001). All of the age groups synchronised their behaviour to a much greater extent than expected by chance. The sucklers showed a higher degree of synchrony of activity (K; 0.890 versus 0.776 versus 0.682 for sucklers, weaners and growers respectively, S.E.D. 0.013, p < 0.001) and inactivity (K; 0.963 versus 0.905 versus 0.909 respectively, S.E.D. 0.007, p < 0.001), but lower degree of synchrony for object directed behaviour (K; 0.149 versus 0.375 versus 0.370 respectively, S.E.D. 0.025, p < 0.001), than the weaners and growers. The degree of synchronisation of object directed behaviour decreased over the 5-day period, irrespective of age (K; 0.382 versus 0.214 for days 1 and 5 respectively, S.E.D. 0.020, p < 0.001). Significant correlations (r = 0.678–0.879) were found between the degree of synchrony and extent of object interaction only for the sucklers. Since pigs showed behavioural synchronisation, object availability should be considered when providing desirable enrichment in order to avoid excessive competition in larger commercial groups.     

Stage-synchronized seminiferous epithelium in rats after manipulation of retinol levels.

Optimal conditions for obtaining stage-synchronization of the seminiferous epithelium were investigated. In this study, 147 rats were subjected to protocols in which vitamin A deficiency was induced by feeding a diet without retinol (R-ol) or retinoic acid (RA), followed by maintenance on a diet containing RA and supplementation of R-ol by injection and diet. An acceptable degree of stage synchronization and recovery of the seminiferous epithelium was observed in 90 (61%) of the 147 rats. The effects on synchrony of variations in the protocol, including the degree of deficiency before RA maintenance, the dose and duration of RA maintenance, and the manner of injection of R-ol, were tested. Initiation of maintenance on RA when a medium degree of deficiency was achieved (4-12 g of weight loss, 3-6 days without growth) resulted in a more reliable (80% of the rats) induction of synchrony than did initiation of maintenance on RA at either a less (70% synchronized rats) or more severe (50-60% synchronized rats) deficiency. Maintenance on food containing 10 mg/kg RA gave better and more reliable synchrony (70%) than maintenance on food containing 5 mg/kg RA (less than 40%). Although the duration of this maintenance did not influence the degree of synchrony, the reliability was lower when maintenance was continued for a month or more (54%). During the interval from 33 to 128 days after resupplementation, the degree of synchronization decreased, as did the predictability of the stages, while the restoration of spermatogenesis increased. Linear regression, performed on the location of the median point of synchronization, indicated that spermatogenesis progressed at a rate of 12.4 days per cycle. The median stage of synchronization, predicted by this regression line, differed by an average of 8% of the cycle from the actual location in individual rats. Extrapolation of the regression line indicated that spermatogenesis was reinitiated in mid-to-late stage VII.     

Anomalous Cases of Conjugation and Spontaneous Autogamy in Stylonychia mytilus: A Note on the Asexuality of Early Conjugants

Four types of anomalous conjugation were documented in Stylonychia mytilus. Type I pairs were formed between mates of different sizes. These pairs exhibited an abnormal site of fusion in at least one of the mates, and the mates might face each other ventrally throughout conjugation instead of the normal side-by-side position. Type I pairs underwent sexual nuclear development and proceeded with the first cortical reorganization as in normal conjugants. Type II involved pairing at the anterior ends of mates with ventral surfaces facing the same direction. These pairs also underwent sexual nuclear development. Hence, aberrant orientation of the mates, and also ectopic sites of cytoplasmic fusion, if extensive, would permit sexual development. Type III pairs were united ventral-to-ventral with their anterior-left sides at the adoral zone of membranelles, and remained as such throughout conjugation. In these pairs, nuclear and cortical events were typical of the asexual development of physiological reorganization. In Type IV pairs, one mate of the pair possessed a fission furrow and developed two sets of ciliature typical of binary fission, while the other mate might undergo physiological reorganization or binary fission. Type III and Type IV pairs thus reveal the asexual state of early conjugants, which can pursue either one of the two modes of asexual cortical reorganization; these cases reinforce the notion of overlap of asexual and sexual cycles during conjugation of hypotrichs. Spontaneous autogamy was documented for the first time for this genus. The autogamonts proceeded with nuclear development and with the first cortical reorganization. Some probably underwent second and third reorganizations, as in conjugants, but accompanied by abnormalities, particularly in the stages beyond fertilization. Post-autogamous clones were nonviable except for one dubious case.