Competitive habitat utilization in the damselfly,Mnais nawai (Zygoptera: Calopterygidae) coexisting with a related species,Mnais pruinosa

Reproductive behaviors related to habitat utilization were studied in males of the damsefly,Mnais nawai, which has two male forms, territorial orange-winged males (nawai) and non-territorial pale-orange-winged males (sahoi), at the upper part of a mountain stream where they partiallycoexist with a related species,Mnais pruinosa, which also has two male forms, territorial orange-winged males (esakii) and non-territorial hyaline-winged males (strigata). These two species showed parapatric distribution; the lower part of the stream was occupied byM. nawai, and the upper part byM. pruinosa. In the present study, cross-matings occurred between bothMnais species, although normal intraspecific matings occurred more frequently than cross-matings. Territorial males of both species copulated with conspecific females that entered their territory and guarded the ovipositing females, probably to avoid sperm displacement resulting from subsequent copulations. Severe competition for oviposition sites by territorial males even occurred between the two species. On the other hand, non-territorial males of both species have alternative mating strategies (including several tactics such as sneaking, takeover and interception). The possible benefits from conflict among territorial males of both species is discussed.     

Synchronization of reproductive period among the two male forms and female of the damselflyMnais pruinosa selys (Zygoptera: Calopterygidae)

Ecological parameters in a population ofMnais pruinosa were investigated in a mountain stream. In the study area, there were two forms of male with regard to wing color, the orange-winged male (esakii) and the hyaline-winged male (strigata), and only one female form with hyaline wings. Emergence of adults began in late April, and the flying season ended in late June. The time after emergence was spent in maturation, and the insects began to mate when they reached maturity. Longevity of adults was 17.6 days foresakii males, 18.4 days forstrigata males and 21.9 days for females. There was little difference in emergence time, maturation period, survivorship curve and longevity among the two male forms and female. In other words, the period for reproductive activities was perfectly coincident among them. The factors influencing the synchronization of emergence were discussed.     

Age-Specific Behavior and Habitat Selection of Adult Male Damselflies, Calopteryx maculata (Odonata: Calopterygidae)

We compared the age, movement, and time–activity budgets of male Calopteryx maculata damselflies occupying off-stream tree-fall gaps with those at stream sites within a 10-ha woodland. All males collected at off-stream sites were younger than males collected at stream sites—as indicated by their significantly higher wing transmittance. Thirty-three percent of teneral males marked at off-stream gaps moved to stream sites within 4 days (mean distance = 140 m), while mature males marked at stream sites never left the stream. In contrast to stream site males, off-stream males spent significantly more time capturing prey and never engaged in aggressive interactions with other males. Behavioral differences were not due to variations in the operative body temperature. However, malaise trapping revealed a greater frequency of suitable prey in forest light gaps. Our findings support the idea that teneral male Calopteryx leave their emergence sites along the stream for off-stream light gaps to forage without interference and build the energy reserves necessary to attain and hold streamside territories.     

Reproductive behavior of females and its relation to the mating success of two male forms inMnais pruinosa (Zygoptera: Calopterygidae)

The behavior of females of the damselflyMnais pruinosa was observed in the breeding season. Males consisted of both territorial (esakii) and non-territorial (strigata) types in the study area. Females sometimes arrived at a stream for mating and oviposition. The staying time of females in the stream was 1–3 hours, while that of males was 5–8 hours. The proportion of the males that copulated with the females did not differ between the two male forms present in the stream. In the Calopterygidae, of which the females usually perform multiple copulation, oviposition without subsequent recopulation is considered to be advantageous for a mated male in order to avoid the risk of sperm displacement. From such a viewpoint, the time spent for oviposition was measured for females that arrived at the stream after copulation with different male forms. The ratio was 69.1: 11.5–30.9 betweenesakii andstrigata males. Assuming that the oviposition time is proportional to the number of fertilized eggs laid, this ratio would represent the relative reproductive success of the two male forms. In fact, the relative abundance of the two male forms was also biased in favor ofesakii males (61.7:38.3) in this population. The mechanism of coexistence of the two male forms is discussed in relation to their reproductive success.     

Differences in adult and reproductive lifespan in the two male forms ofMnais pruinosa costalis selys (Odonata: Calopterygidae)

The damselflyMnais pruinosa costalis (Odonata: Calopterygidae) is unusual in that males are dimorphic, existing as clear-winged non-territorial ‘sneaks’ and orange-winged territorial ‘fighters’. Here we report the results of population census data and behavioural observations in the field and laboratory, and present estimates of emergence period, reproductive period, total lifespan, and reproductive success of each morph. Clear-winged males are smaller and have lower daily reproductive success than orange-winged males, but live for longer in the field and laboratory. We accounted for the difference in the ‘operational reproductive life’ of the two morphs and estimated lifetime reproductive success: there was no difference between clear-winged and orange-winged males. We discuss possible mechanisms for the maintenance of the two forms.     

Male mate choice and the potential for complex mating dynamics in the tree lizard (Urosaurus ornatus)

A growing body of literature is recognizing that males may also play a role in the mating process by behaving non‐randomly toward potential female mates during courtship. In numerous species, discrete color polymorphisms in males are inferred to represent alternative mating tactics, which often correspond with concomitant asymmetries in ecology and behavior. In terms of their mating behavior, these ecological outcomes of a color polymorphism should affect a morph's likelihood and frequency of encountering females in a population, possibly favoring the evolution of morph‐specific mating preferences. Knowledge of how male morphs contribute to a species’ overall mating dynamics will improve our understanding of how sexual selection shapes phenotypic diversity in color polymorphic systems. We conducted a mate choice experiment to evaluate the extent and morph specificity of non‐random mating preferences by male ornate tree lizards, Urosaurus ornatus. We observed the behavior of blue and yellow males in an experimental arena in response to a choice between an orange or yellow female. We found that blue males preferred yellow females over orange females, and although yellow males visited females more often than blue males overall, their attention was not morph‐specific. Given male morph differences in choosiness, and their differences in social dominance, we conclude that female throat color may be partly under sexual selection in U. ornatus. However, a lack of concordance between male and female mating preferences (drawn from an earlier study) suggests that overall mating dynamics may serve to maintain, rather than enhance, color morph differences in this species.     

Description of the Final Stadium Larva of Hetaerina rosea Selys (Zygoptera: Calopterygidae)

The final stadium larva of Hetaerina rosea is here described and illustrated for the first time based on specimens collected in Corrientes and Buenos Aires provinces, Argentina.     

Asymmetric intraspecific competition among larvae of the damselfly Ischnura elegans (Zygoptera: Coenagrionidae)

Abstract. 1. A laboratory competition experiment is described in which the growth and development rates of larvae of the damselfly Ischnura elegans (Lind.) were measured over an entire instar.
2. Two larval instars which commonly occur together in the field were used in the experiment; they were maintained with a superabundance of prey and either larvae from the same or the larger/smaller instar.
3. Small larvae suffered increased development times and decreased size increases at the moult in the presence of large larvae but similar interference effects were not evident when these smaller larvae were in the presence of other small larvae.
4. Development time and size increases of large larvae were not significantly affected by the presence of small larvae.
5. Irrespective of the instar combinations investigated, interference effects were reduced when there were more perches available, although in only a few cases was this reduction significant.
6. The consequences of the asymmetric competition reported in the experiment for the study of lifetime reproductive success in damselflies are discussed. Late emerging adults may incur reduced reproductive success.     

Remarkable iridescence in the hindwings of the damselfly Neurobasis chinensis chinensis (Linnaeus) (Zygoptera: Calopterygidae)

The bright green dorsal iridescence of the hindwings of Neurobasis chinensis chinensis males, very rare in Odonata, is known to play a significant role in their courtship behaviour. The mechanism responsible for such high contrast and spectrally pure colour has been investigated and found to be optical interference, producing structural colour from distinct laminations in the wing membrane cuticle. The ventral sides of these iridescent wings are dark brown in colour. In a single continuous membrane of wing cuticle, this is an effect that requires a specialized structure. It is accomplished through the presence of high optical absorption (kappa = 0.13) within two thick layers near the ventral surface of the wing, which leads to superior dorsal colour characteristics. By simultaneously fitting five sets of optical reflectivity and transmissivity spectra to theory, we were able to extract very accurate values of the complex refractive index for all three layer types present in the wing. The real parts of these are n = 1.47, 1.68 and 1.74. Although there is often similarly significant dorsal and ventral colour contrast in other structurally coloured natural systems, very few system designs comprise only a single continuous membrane.     

Pre-oviposition mate-guarding and mating behaviour of Argia vivida (Odonata: Coenagrionidae)

Abstract. 1. At Halcyon Hotsprings, British Columbia, Canada, male and female Argia vivida Hagen encountered to mate in two different ways.
2. In the morning (before 12.30 hours solar time), males basked at sunspots in the forest and darted out at passing females, attempting to take them in tandem (the first method of encounter).
3. If a male was successful, the pair engaged in a 31.3±4.8 min copulation followed by an hour of tandem flight before beginning oviposition.
4. As the day progressed, unmated males moved slowly toward the water and arrived at the water at about the same time as the earliest ovipositing pairs (1131±27.5 min solar time).
5. Males retained their grasp on their mates during oviposition (contact-guarding) but since some tandems separated during oviposition, non-tandem males at the water could capture recently released, gravid females (the second method of encounter).
6. The new pairs performed a brief copulation (10.2±3.38 min) and began ovipositing immediately thereafter.
7. Some females that avoided recapture attempted to oviposit unguarded.
8. We believe the long duration of morning copulations and period of tandem constitute a male strategy, which we call 'pre-oviposition guarding', to guard females until it is warm enough at the oviposition site for the females to begin ovipositing.
9. Separation of tandems during oviposition may be initiated by either member of the pair and we suggest that one benefit to a female of leaving a guarding mate is increased efficiency of oviposition when the intensity of male harassment is low.
10. The mating system of A. vivida thus comprises a series of complementary male and female mating behaviours.     

Responses of rectal pumping to oxygen lack by larval Calopteryx splendens (Zygoptera: Odonata)

Abstract. Larvae of most stadia of Calopteryx splendens (Harris) have been shown to pump water in and out of the rectum for much of the time. In each cycle of pumping there are from three to seven inhalant stokes which progressively fill the ileum with water. A short pause intervenes before a strong compression of segments 5–8 expels the water in a jet. The pattern resembles the gulping ventilation of aeshnid and libellulid dragonfly larvae. Larvae made hypoxic or otherwise stressed show a rise in the frequency and amplitude of pumping. The increase in frequency is brought about by a reduction of the pause duration, while the gain of amplitude results from a rise in both the number of inhalations/cycle and the magnitude of the stroke volume of each inhalation. Some evidence concerning the role of stretch receptors in the execution of the cycle is presented. It is proposed that rectal pumping promotes the exchange of ions and dissolved gases between the rectal water and the hindgut epithelia, supplying mainly the chloride-absorbing cells in the rectal pads. Pumping also stirs the water in the vicinity of the caudal lamellae which are probably the insect's major sites of gaseous exchange.     

Using remote sensing image texture to study habitat use patterns: a case study using the polymorphic white-throated sparrow (Zonotrichia albicollis)

Aim Ecological factors that vary along spatial scales can greatly influence the outcome of evolution. However, often it is difficult for researchers to identify significant ecological variables that might be associated with the geographical distribution of phenotypes and genotypes in nature. In this paper, we use remote sensing image texture analyses to investigate breeding territories of the white‐throated sparrow (Zonotrichia albicollis), a polymorphic passerine species that has been shown to segregate spatially according to morph. Our aim was to examine how an integrative measure of landscape feature, such as image texture, could be used to distinguish subtle differences in habitat use and, presumably, habitat selection. Location A population of white‐throated sparrows located 1.5 miles east of the Cranberry Lake Biological Station in the Adirondack Mountains of St Lawrence County, New York, was used in this study; the study site itself was approximately 1 km². Methods We analysed 27 territories (16 white males; 11 tan males) to determine if image texture can be used to distinguish differences in vegetation and structure in territories of the two colour morphs. First (variance)‐ and second (homogeneity)‐order image texture measurements with three different window sizes were derived using the green, red, and near infrared bands of Advanced Spaceborne Thermal Emission and Reflection Radiometer (ASTER) data. Texture was then placed into t‐tests to determine statistical differences between the territories of the two male morphs. Results Significant differences between the territories of the two morphs were found in both variance and homogeneity texture measures in the red and green ASTER bands in all three window sizes. There were no differences with any near infrared texture measures. Main conclusions Examination of first‐ and second‐order statistical measures indicated that the territories of tan males have significantly more spectral variance than the territories of white males, while white males’ territories are significantly more homogeneous. These findings are consistent with the previous finding of the morphs settling in ‘high’ and ‘low’ neighbour density habitats that differ with respect to the amount of useable area on their borders. Previous study has also suggested that white and tan male territories differ according to overall landscape structure, not vegetation composition. Therefore, we suggest that red and green spectra may be more useful when examining differences in territory structure and possibly territory quality of similar edge‐type species. Our results suggest that image texture is a useful tool for ecological and evolutionary modelling of the distributions of phenotypes/genotypes across landscapes.     

Effects of delayed mating and male mating history on longevity and reproductive performance of the rice stem borer, Chilo suppressalis (Walker) (Lep., Pyralidae)

Abstract:  The mating potential, effects of delayed mating and male mating history on longevity and reproductive performance of female rice stem borer (RSB), Chilo suppressalis , were investigated under laboratory conditions. Given the opportunity, RSB males copulated an average of 2.7 times, while females generally mated only once. Females were more severely affected by mating delay than males in terms of female longevity and reproductive performance. With increasing age at mating, females' longevity increased, while oviposition period, fecundity and egg fertility decreased. Mean fecundity and egg fertility of females mated 1 and 7 days after emergence were 251.3% and 99.2%, and 96.2% and 75.5% respectively. Both were reduced significantly when female mating was delayed beyond 4 days after emergence. However, irrespective of the first mating age and different mating history of the males, the results indicated that both did not result in a significant difference in the quality of their contributions to female longevity and reproductive performance. The results obtained in this study are discussed in relation to the potential effect on pheromone control of RSB.     

Shifts in reproductive assurance strategies and inbreeding costs associated with habitat fragmentation in Central American mahogany

The influence of habitat fragmentation on mating patterns and progeny fitness in trees is critical for understanding the long-term impact of contemporary landscape change on the sustainability of biodiversity. We examined the relationship between mating patterns, using microsatellites, and fitness of progeny, in a common garden trial, for the insect-pollinated big-leaf mahogany, Swietenia macrophylla King, sourced from forests and isolated trees in 16 populations across Central America. As expected, isolated trees had disrupted mating patterns and reduced fitness. However, for dry provenances, fitness was negatively related to correlated paternity, while for mesic provenances, fitness was correlated positively with outcrossing rate and negatively with correlated paternity. Poorer performance of mesic provenances is likely because of reduced effective pollen donor density due to poorer environmental suitability and greater disturbance history. Our results demonstrate a differential shift in reproductive assurance and inbreeding costs in mahogany, driven by exploitation history and contemporary landscape context.     

Early sexual maturity in male hamadryas baboons (Papio hamadryas hamadryas) and its reproductive implications

We present data on sexual maturity in young hamadryas baboon males (Papio hamadryas hamadryas) and its reproductive consequences in a large captive baboon colony. Hamadryas baboons live in a multilevel social system, with one-male units (OMUs) as the smallest social entity. Male leaders of OMUs are believed to monopolize matings within their OMUs; hence mating is believed to be polygynous and monandrous. In a captive colony of hamadryas baboons, we found evidence that young males less than 4 years old fathered at least 2.5% of 121 offspring born subsequent to vasectomy of all adult males, and males aged 4-5 years fathered at least 16.5% of the offspring. Additional evidence that these young males are able to sire offspring came from a morphological comparison of sperm from hamadryas males of different ages. The sperm of a 48-month-old hamadryas baboon were morphologically indistinguishable from viable sperm from adult males, whereas sperm from a 45-month-old male showed some aberrations. If successful copulations by adolescent males constitute a regular pattern even in free-ranging hamadryas baboons, a hamadryas male's chances to reproduce would not be limited to his role as an OMU leader as previously assumed, and a male's reproductive career would consist of two phases: the adolescent phase, and the OMU leader male phase.     

Scale effects of habitat selection in breeding Nuthatches (Sitta europaea) in two different woodlands

Summary Observations over a 14 year study revealed that Nuthatches had significantly higher breeding densities in a deciduous woodland than in coniferous-mixed woodland. However, the variability in breeding densities between years correlated in the two woodlands. In selecting a breeding site, birds may have to decide on different spatial scales. Possibly, these scales are reflected in a hierarchical series of decisions that might differ according to habitat quality. We correlated the frequency of nest box occupation by Nuthatches over 14 years with habitat parameters and PCA scores of different spatial scales (roughly 0.1 ha and 1 ha). In the coniferous-mixed woodland Nuthatches preferred similar habitat parameters at the small and the large scale like oaks, open areas and forest edge. In the deciduous woodland, however, a marked difference in habitat preference between small and large scale was shown: on the small scale parameters directly associated with nest box characteristics were significantly preferred. On the large scale only structural and floristic parameters associated with the general quality of the woodland were important for breeding site selection. Here tree stands with a superabundance of pine trees were avoided and the abundant oak trees were not important for the selection of a nest box. By further subdividing the very heterogeneous deciduous woodland into two parts, a stand of young trees and the rest of the woodland, opposing habitat choices could be shown. Thus scale effects of habitat choice are dependent upon habitat quality and heterogeneity.

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Einfluß des räumlichen Maßstabs auf die Habitatwahl von Kleibern (Sitta europaea) zur Brutzeit in zwei unterschiedlichen Waldgebieten

Zusammenfassung In einer Langzeituntersuchung über 14 Jahre wurde die Nistkastenbelegung durch Kleiber in zwei unterschiedlichen Waldgebieten, einem Laubwald und einem Nadel-Laub-Mischwald, untersucht. Im Laubwald waren 80 Nistkästen, im Nadel-Laub-Mischwald 92 Nistkästen in 30 × 30 m Rastern aufgehängt. In jedem dieser Rasterquadrate wurden 18 Habitatparameter aufgenommen. Beide Waldgebiete unterschieden sich in den meisten Parametern signifikant. Die Brutpaardichten waren im Laubwald signifikant höher als im Nadel-Laub-Mischwald. Jedoch korrelierte die Variabilität der Brutpaardichten zwischen den Untersuchungszeiträumen in beiden Gebieten. Nistplatzwahl wurde in zwei räumlichen Maßstäben untersucht. Der kleine Maßstab umfaßte die Fläche von 30 × 30 m um jeden Nistkasten (ca. 0,1 ha), der große Maßstab hatte die Fläche von ca. 1 ha (9 Nistkästen, 90 × 90 m). Auf dem kleinen Maßstab wurde die Nistkastenbelegung für jeden Nistkasten über den Untersuchungszeitraum aufsummiert. Auf dem großen Maßstab wurde für jeden Nistkasten und die ihn umgebenden 8 Nistkästen die Belegung gemittelt. Dadurch ergaben sich für großen und kleinen Maßstab identische Stichprobengrößen. Die so ermittelten Nistkastenbelegungen wurden mit den Habitatparametern und entsprechenden Hauptkomponentenfaktoren korreliert.Im Nadel-Laub-Mischwald bevorzugten Kleiber bei beiden Maßstäben Eichen, offene Flächen und Waldränder. Im Laubwald unterschied sich die Habitatpräferenz zwischen den Maßstäben: bei kleinem Maßstab wurden Nistkasten-Parameter (wie z. B. die Höhe) signifikant bevorzugt. Bei großem Maßstab hingegen waren nur Strukturparameter sowie Pflanzenarten für die Nistplatzwahl entscheidend. Waldbereiche mit einem Überangebot an Kiefern wurden gemieden, die häufig und gleichmäßig vorkommenden Eichen hatten bei der Nistplatzwahl keine Bedeutung. Bei weiterer Unterteilung des sehr heterogenen Laubwaldes in zwei Einheiten, einen jungen Baumbestand und den Rest des Waldes, wurde gegensätzliche Habitatwahl nachgewiesen. Daraus folgt, daß bei der Habitatwahl durch unterschiedliche Maßstäbe verursachte Effekte von Habitatqualität und -heterogenität abhängen.

     

Emergence of the damselfly Pyrrhosoma nymphula (Sulzer) (Zygoptera: Coenagrionidae) from two adjacent ponds in northern England

Emergence of the damselfly Pyrrhosoma nymphula was followed closely at two similar and adjacent ponds in northern England. Males emerged earlier than females at both ponds (significantly so at one). The sex ratio was significantly male biased at both ponds. Size of emerging adults declined through the emergence period at both ponds. Adults emerging from one pond did so significantly earlier and were significantly larger than those from the other pond. The density of larvae was approximately twice as high in the pond from which larvae emerged early; this pond was also slightly deeper and usually achieved higher maximum daily water temperatures. These findings are discussed in the light of the conventional view of seasonal regulation in a spring-emerging damselfly.     

Starvation and the Mating Success of Wild Male Mediterranean Fruit Flies (Diptera: Tephritidae)

Recent evidence suggests that the nutritional state of male Mediterranean fruit flies, Ceratitis capitata (Wied.) (medfly), is an important influence on various components of their reproductive biology, including mating success. The objective of the present study was to examine experimentally the effect of temporary starvation on the mating success of wild male C. capitata. Males were maintained on protein–sugar or sugar-only diets, and for each diet we compared the mating success of continuously fed males versus males starved for 18 or 24 h immediately before testing. In trials conducted on field-caged, host trees, males starved for 24 h obtained only about half as many matings as fed males for both diets. However, when the starvation period was 18 h, starved males reared on the protein–sugar diet mated significantly less frequently than fed males, whereas starved males reared on sugar mated as often as fed males. Measurements of male pheromone calling and female attraction revealed that reduced mating success likely reflected the decreased signaling activity of starved males.     

The effects of temperature and prey density on the development rates and growth of damselfly larvae (Odonata: Zygoptera)

Abstract. 1. The effects of prey density and temperature on the feeding and development rates of several late instars of the larvae of three common European damselflies ( Lestes sponsa (Hansemann), Coenagrion puella (L.) and Ischnura elegans van der Linden) were investigated in a laboratory experiment.
2. Functional responses were used to estimate maximum feeding rates. Maximum feeding rates were compared between species, instars and temperatures by expressing prey consumption in terms of prey biomass consumed per unit predator biomass. Lestes was capable of feeding at almost twice the rate of either Coenagrion or Ischnura.
3. Higher feeding rates led to faster development rates and there was an interaction between species and temperature. With the exception of those with very low feeding rates, larvae maintained at higher temperatures, but similar feeding rates, developed faster. Under similar conditions of temperature and feeding rate, Lestes larvae developed faster than larvae of either Coenagrion or Ischnura.
4. Faster development rates at similar rates of food intake were achieved at the cost of reduced size-increases between instars.
5. The differences in the responses of the three species are discussed in the light of their respective life histories, and with reference to a recent model of population regulation in damselflies.     

Nearshore fish habitat utilization and species associations in Lake Opinicon (Ontario,Canada)

Synopsis The bulk of Lake Opinicon fish biomass is concentrated in the physically diverse inshore areas. Quantitative analysis of the community compositions of the various inshore habitat types (weedy inlets, sandy shallows, rock shelf, gravel, etc.) showed that each supported a characteristic assemblage of fishes with a relatively constant species and year class composition. This was maintained throughout the season despite a drop in fish biomass in late summer when there was a progressive movement into the offshore waters.Weedbed areas supported the highest biomass, and greatest species and year class diversity. Thereafter, in declining order of richness, were rocky and sandy areas. A few fish species were restricted to single habitat types, e.g.Notropis heterodon to inshore weedbeds.Lepomis macrochirus, the commonest species in the lake was, by contrast, versatile and occurred in all inshore habitats. Commonly the numbers of a species in a habitat differed between day and night. Significant diel movements between habitats characterized two nocturnal feedersPomoxis nigromaculatus andIctalurus nebulosus.