Use of statistical models based on radiographic measurements to predict oviposition date and clutch size in rock iguanas (Cyclura nubila)

The ability to noninvasively estimate clutch size and predict oviposition date in reptiles can be useful not only to veterinary clinicians but also to managers of captive collections and field researchers. Measurements of egg size and shape, as well as position of the clutch within the coelomic cavity, were taken from diagnostic radiographs of 20 female Cuban rock iguanas, Cyclura nubila, 81 to 18 days prior to laying. Combined with data on maternal body size, these variables were entered into multiple regression models to predict clutch size and timing of egg laying. The model for clutch size was accurate to 0.53 ± 0.08 eggs, while the model for oviposition date was accurate to 6.22 ± 0.81 days. Equations were generated that should be applicable to this and other large Cyclura species. © 1995 Wiley-Liss, Inc.     

Reproductive dynamics of a tropical freshwater crocodilian: Morelet's crocodile in northern Belize

Morelet's crocodile Crocodylus moreletii has not been well-studied and many aspects of its life history are unknown. In particular there is a notable paucity of information on nesting and reproductive ecology. We studied the nesting ecology of Morelet's crocodile in northern Belize from 1992 through 1995. Nesting occurs at the onset of the wet season in mid-June and continues through mid-July (mean oviposition date =1 July±10 days). Eggs hatch from mid-August through mid- to late September. Nesting effort at our primary study site remained relatively constant during 1992, 1993 and 1995, but nearly doubled in 1994; this appeared to reflect a regional trend. Natural and man-made islands are heavily used as nesting sites. Nesting success in 1993 and 1994 was consistently higher on natural islands when compared with man-made islands or shoreline sites. Nest losses were primarily due to flooding and raccoon Procyon lotor predation. Losses from predation were greatest in 1994 when unseasonably low water levels facilitated predator access to nests. Females probably reach sexual maturity in 7–8 years after attaining a total length of 150 cm. Mean clutch size (25.0±7.6; range=9–42; n =73) did not differ among years. Mean clutch size, egg width (EW), egg length, egg mass (EM) and clutch mass were positively correlated with female snout–vent length (SVL). Mean EW was the best predictor of female SVL. A partial correlation analysis of egg and clutch attributes found that independent of female SVL, EM increases with increasing clutch size.     

Nesting ecology of a naturalized population of Mallards Anas platyrhynchos in New Zealand

Investigating the reproductive ecology of naturalized species provides insights into the role of the source population's characteristics vs. post‐release adaptation that influence the success of introduction programmes. Introduced and naturalized Mallards Anas platyrhynchos are widely established in New Zealand (NZ), but little is known regarding their reproductive ecology. We evaluated the nesting ecology of female Mallards at two study sites in NZ (Southland and Waikato) in 2014–15. We radiotagged 241 pre‐breeding females with abdominal‐implant transmitters and measured breeding incidence, nesting chronology and re‐nesting propensity. We monitored 271 nests to evaluate nest survival, clutch and egg size, egg hatchability and partial clutch depredation. Breeding incidence averaged (mean ± se) 0.91 ± 0.03, clutch size averaged 9.9 ± 0.1 eggs, 94 ± 2% of eggs hatched in successful nests, partial depredation affected 6 ± 1% of eggs in clutches that were not fully destroyed by predators, and re‐nesting propensity following failure of nests or broods was 0.50 ± 0.003. Nesting season (first nest initiated to last nest hatched) lasted 4.5 months and mean initiation date of first detected nest attempts was 28 August ± 3.3 days. Smaller females were less likely to nest, but older, larger or better condition females nested earlier, re‐nested more often and laid larger clutches than did younger, smaller or poorer condition females. Younger females in Southland had higher nest survival; cumulative nest survival ranged from 0.25 ± 0.007 for adult females in Waikato to 0.50 ± 0.007 for yearling females in Southland. Compared with Mallards in their native range, the nesting season in NZ was longer, clutches and eggs were larger, and nest survival was generally greater. Different predators and climate, introgression with native heterospecifics and/or the sedentary nature of Mallards in NZ may have contributed to these differences.     

Geographic variation in reproductive traits and trade-offs between size and number of eggs of the Chinese cobra (Naja atra)

We collected gravid Chinese cobras (Naja atra) from one island (Dinghai) and three mainland (Yiwu, Lishui and Quanzhou) populations in south‐eastern China to study geographical variation in female reproductive traits and the trade‐off between the size and number of eggs. We then conducted an common experiment on cobras from two of the four populations to further identify factors contributing to the observed trade‐offs. The mean size (snout–vent length) of the smallest five reproductive females increased with increasing latitude. Oviposition occurred between late June and early August, with females from the warmer localities laying eggs earlier than those from the colder localities. Maternal size was a major determinant of the reproductive investment in all populations, with larger females producing not only more but also larger eggs. Clutch size was more variable than egg size within and among populations. The observed geographical variation in clutch size, egg size, clutch mass and post‐oviposition body condition was not a simple consequence of variation in maternal size among populations, because interpopulation differences in these traits were still evident when the influence of maternal size was removed. The upper limit to reproductive investment was more likely to be set by the space availability in the island population, but by the resource availability in the three mainland populations. Trade‐offs between size and number of eggs were detected in all populations, with females that had larger clutches for their size having smaller eggs. Egg size at any given level of relative fecundity differed among populations, primarily because of interpopulation differences in the resource availability rather than the space availability. Except for the timing date of oviposition and the mean size of the smallest five reproductive females, all other examined traits did not vary in a geographically continuous trend. The common garden experiment, which standardized environmental factors, synchronized the timing date of oviposition, but it did not modify the conclusion drawn from the gravid females collected from the field. The observed geographical variation in the female reproductive traits could be attributed to the consequence of the effects of either proximate or ultimate factors. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 85 , 27–40.     

Incubation capacity contributes to constraints on maximal clutch size in Brent Geese Branta bernicla nigricans

Lack ( 1967 ) proposed that clutch size in species with precocial young was determined by nutrients available to females at the time of egg formation; since then others have suggested that regulation of clutch size in these species may be more complex. We tested whether incubation limitation contributes to ultimate constraints on maximal clutch size in Black Brent Geese (Black Brant) Branta bernicla nigricans. Specifically, we investigated the relationship between clutch size and duration of the nesting period (i.e. days between nest initiation and the first pipped egg) and the number of goslings leaving the nest. We used experimental clutch manipulations to assess these questions because they allowed us to create clutches that were larger than the typical maximum of five eggs in this species. We found that the per‐capita probability of egg success (i.e. the probability an egg hatched and the gosling left the nest) declined from 0.81 for two‐egg clutches to 0.50 for seven‐egg clutches. As a result of declining egg success, clutches containing more than five eggs produced, at best, only marginally more offspring. Manipulating clutch size at the beginning of incubation had no effect on the duration of the nesting period, but the nesting period increased with the number of eggs a female laid naturally prior to manipulation, from 25.4 days (95% CI 25.1–25.7) for three‐egg clutches to 27.7 days (95% CI 27.3–28.1) for six‐egg clutches. This delay in hatching may result in reduced gosling growth rates due to declining forage quality during the brood rearing period. Our results suggest that the strong right truncation of Brent clutches, which results in few clutches greater than five, is partially explained by the declining incubation capacity of females as clutch size increases and a delay in hatching with each additional egg laid. As a result, females laying clutches with more than five eggs would typically gain little fitness benefit above that associated with a five‐egg clutch.     

Variations in fecundity over catchment scales: Implications for caddisfly populations spanning a thermal gradient

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The use of ultrasonography to assess reproductive investment and output in pythons

Reproductive investment and output are integral fitness components, often incorporated into life‐history trade‐off models and important to population dynamics. The trade‐offs associated with reproduction can be dramatic in species such as snakes that make especially large investments into reproduction. Unfortunately, traditional methods used to determine reproductive investment and output are effective in many (but not all) situations. Thus, we used portable ultrasonography to serially estimate reproductive investment and reproductive output in three python species that exhibit significant variation in phylogeny, geographic range, body size, egg size, and clutch size: ball pythons (Python regius), Children's pythons (Antaresia childreni), and water pythons (Liasis fuscus). At each time point of measurement (range: 1–49 days pre‐oviposition), ultrasound estimates of viable clutch size were highly accurate in all three species. However, ultrasound estimates of mean viable egg mass, and thus viable clutch mass, significantly differed from the actual values (range: 23–73% error). Interestingly, this error was considerably smaller as females approached oviposition, suggesting that female pythons transfer a significant amount of water into their eggs during the week before oviposition. Thus, water balance during late‐stage egg development may be an integral part of reproductive success. The results obtained in the present study form the foundation for future assessments of reproductive investment, and also provide insight into the use of ultrasound technology to assist such efforts. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 772–778.     

Breeding ecology of Red‐faced Cormorants in the Pribilof Islands,Alaska

ABSTRACT Red‐faced Cormorants (Phalacrocorax urile) are North Pacific endemics recognized as a vulnerable species, but little is known about their breeding ecology. We studied Red‐faced Cormorants on St. Paul Island, Alaska, from 1975 to 2009, with more detailed data collected in 2004 and 2005. Mean clutch sizes in 2004 (3.2 ± 0.8 [SD] eggs) and 2005 (3.1 ± 0.8 eggs) were similar to the long‐term average (2.9 ± 0.3 eggs from 1976 to 2009). The mean laying interval in 2004 and 2005 was 2.15 ± 0.80 d (N= 407), and the mean egg period (number of days between laying of an egg and hatching) was 31.1 ± 1.4 d (N= 158). Approximately 64 ± 17% of eggs hatched during the period from 1975 to 2009. The mean number of chicks per nest in 2004 and 2005 was 2.8 ± 0.8 (N= 232), and the mean number of fledglings per initiated nest in all years was 1.22 ± 0.52. Chicks fledged 46 to 66 d posthatching. In 2004 and 2005, the primary causes of egg loss were predation by Arctic foxes (Vulpes lagopus) and destruction of eggs and abandonment of nests due to storms. Starvation was the primary cause of nestling mortality in both years. Because chicks are dependent on parents to provide food for over 45 d, consistent near‐shore foraging opportunities must be available. From 1975 to 2009, Red‐faced Cormorants experienced only 1 yr of complete reproductive failure (1984). The consistent reproductive success of Red‐faced Cormorants suggests that conditions may be relatively stable for this species on St. Paul Island, or that the variability in their breeding ecology (e.g., phenology, clutch sizes, and incubation strategies) provides the flexibility needed to successfully fledge some chicks nearly every year.     

Bionomics of the acarophagous ladybird beetle Stethorus tridens fed Tetranychus evansi

The bionomics of Stethorus tridens Gordon fed Tetranychus evansi Baker & Pritchard were studied in the laboratory. The number of prey consumed by S. tridens increased with increasing instar levels and the total mean number consumed during immature development was 184.1 ± 18.02 T. evansi nymphs per individual. For adult male and adult female, the daily consumption was 41.3 ± 0.80 and 67.8 ± 1.69 nymphs, respectively. Stethorus tridens successfully developed to adulthood between 20 and 30°C but failed at 10, 15 and 35°C. The lower thermal threshold for egg‐to‐adult development estimated via linear regression and the modified Logan model was 9.2 and 8.1°C, respectively. The optimum and maximum temperatures for egg‐to‐adult development were around 29–31 and 32.9°C, respectively. Egg to adult development time was 23.8 ± 0.24, 17.4 ± 0.22, 16.2 ± 0.22 and 12.1 ± 0.16 days at 20, 24, 27 and 30°C, respectively. At 27°C, the sex ratio, expressed as the proportion of females, was 0.54 and the mean preoviposition, oviposition and postoviposition periods were 10.3 ± 0.67, 31.2 ± 4.74 and 30.2 ± 5.24 days, respectively. The oviposition rate was 4.0 ± 0.16 eggs/female/day with a female mean longevity of 71.6 ± 6.19 days and an intrinsic rate of natural increase of 0.104. The potential of S. tridens as a candidate natural enemy of T. evansi is discussed.     

Effects of intraclutch variation in egg size and hatching asynchrony on nestling development and survival in semi‐precocial Herring Gulls

Intraclutch egg size variation may non‐adaptively result from nutritional/energetic constraints acting on laying females or may reflect adaptive differential investment in offspring in relation to laying/hatching order. This variation may contribute to size hierarchies among siblings already established due to hatching asynchrony, and resultant competitive asymmetries often lead to starvation of the weakest nestling within a brood. The costs in terms of chick mortality can be high. However, the extent to which this mortality is egg size‐mediated remains unclear, especially in relation to hatching asynchrony which may operate concomitantly. I assessed effects of egg size and hatching asynchrony on nestling development and survival of Herring Gulls (Larus argentatus), where the smaller size and later hatching of c‐eggs may represent a brood‐reduction strategy. To analyze variation in egg size, I recorded the laying order and laying date of 870 eggs in 290 three‐egg clutches over a 3‐yr period (2010–2012). I measured hatchlings and monitored growth and survival of 130 chicks from enclosed nests in 2011 and 2012. The negative effect of laying date (β = ?0.18 ± SE 0.06, P = 0.002) on c‐egg size possibly reflected the fact that late breeders were either low quality or inexperienced females. The mass, size, and condition of hatchling Herring Gulls were positively related to egg size (all P < 0.0001). C‐chicks suffered from increased mortality risk during the first 12 d, identified as the brood‐reduction period in my study population. Although intraclutch variation in egg size was not directly related to patterns of chick mortality, I found that smaller relative egg size interactively increased differences in relative body condition of nestlings, primarily brought about by the degree of hatching asynchrony during this brood‐reduction period. Thus, the value of relatively small c‐eggs in Herring Gulls may lie in reinforcing brood reduction through effects on nestling body condition. A reproductive strategy Herring Gulls might have adopted to maintain a three‐egg clutch, but that also enables them to adjust the number of chicks they rear relative to the prevailing environmental conditions and to their own condition during the nestling stage.     

On the reproductive biology of the herbivorous spiny-tailed agamid Uromastyx philbyi in western Saudi Arabia

Abstract

The reproductive biology of the Spiny-tailed Agamid Uromastyx philbyi, a herbivorous desert lizard, was studied in western Saudi Arabia. Reproduction is seasonal, with mating in early spring (March), oviposition in late spring (May–June), and hatching in summer (July). The mean clutch size was 6.67 eggs. Eggs are large (mean mass = 7.48 g) and relative clutch mass (RCM) averaged 0.49. Clutch mass and RCM increased with increasing egg mass. Clutch size, egg size, clutch mass, and RCM were significantly correlated with maternal body size.     

Geographical variation in reproductive traits and trade‐offs between size and number of eggs in the king ratsnake,Elaphe carinata

We collected gravid king ratsnakes (Elaphe carinata) from three geographically separated populations in Chenzhou (CZ), Lishui (LS) and Dinghai (DH) of China to study the geographical variation in female reproductive traits and trade‐offs between the size and number of eggs. Not all reproductive traits varied among the three populations. Of the traits examined, five (egg‐laying date, post‐oviposition body mass, clutch size, egg mass and egg width) differed among the three populations. The egg‐laying date, ranging from late June to early August, varied among populations in a geographically continuous trend, with females at the most northern latitude (DH) laying eggs latest, and females at the most southern latitude (CZ) laying eggs earliest. Such a trend was less evident or even absent in the other traits that differed among the three populations. CZ and DH females, although separated by a distance of approximately 1100 km as the crow flies, were similar to each other in most traits examined. LS females were distinguished from CZ and DH females by the fact that they laid a greater number of eggs, but these were smaller. The egg size–number trade‐off was evident in each of the three populations and, at a given level of relative fecundity, egg mass was significantly greater in the DH population than in the LS population. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 701–709.     

Oviposition behavior and progeny allocation of the polyembryonic waspCopidosoma floridanum (Hymenoptera: Encyrtidae)

Oviposition behavior was used to determine the primary clutch size and sex ratio of the polyembryonic wasp Copidosoma floridanumAshmead (Hymenoptera: Encyrtidae) parasitizing Pseudoplusia includens(Walker) (Lepidoptera: Noctuidae). The laying of a female egg was associated with a pause in abdominal contractions during oviposition, while the laying of a male egg was associated with uninterrupted abdominal contractions. Although unmated females produced only male broods, they also displayed male and female egg oviposition movements. Wasps always laid a primary clutch of one or two eggs. For mated females if only one egg was laid, the emerging secondary clutch was all male or female, but if two eggs were laid a mixed brood of males and females was almost always produced. The secondary clutch of single sex broods was usually between 1000 and 1200 individuals, but the secondary clutch of mixed broods averaged 1143 females and 49 males. Thus, the primary sex ratio for mixed broods was 0.5 (frequency males), but the secondary sex ratio was 0.042. Manipulation of the sequence of male and female egg oviposition or of the primary clutch did not produce major alterations in the secondary clutch size or sex ratio.     

On a Poorly Known Rookery of Green Turtles (<Emphasis Type="Italic">Chelonia mydas</Emphasis>) Nesting at the Chabahar Beach,Northeastern Gulf of Oman

There is a poorly known rookery of green turtles (Chelonia mydas) nesting in sandy beaches of Chabahar town, northeastern Gulf of Oman, Iran.This study has been carried out to evaluate nesting activity of this small rookery in 2014 nesting season (June to October). In this study, total clutches were collected and transferred to an artificial hatchery. The peak of nesting occurred from the third parts of August to the end of September. Mean CCL was 106.3 ± 6 cm and mean CCW was 94.5 ± 5 cm. Females laid on average of 99.42 ± 47.8 eggs per clutch. The mean inter-nesting interval was 18.5 days. The observed clutch frequency was 3.4. Mean hatching success was 36.63 ± 4.1%. The incubation period was 61.07 ± 5.4 days. Nest status evaluation represented that major causes for the failure was unhatched egg with no obvious embryo followed by unhatched eggs with obvious but undeveloped embryos > unhatched eggs with developed embryos > hatched eggs but dead embryos. The results achieved in this study are a valuable contribution to cognition of the reproductive ecology of the green turtle population globally and regionally.     

Egg-size variation in the bluethroat (Luscinia s. svecica): constraints and adaptation

We examined inter- and intra-clutch egg-size variation in the bluethroat (Luscinia s. svecica), an open-nesting passerine breeding in the sub-alpine region in southern Norway. By removing first clutches shortly after egg-laying, we induced laying of a repeat clutch. Females significantly reduced the number of eggs from the first to the second nesting attempt, but increased mean egg size. Females in good condition laid significantly larger eggs than those in poor condition. Consistent with predictions of the brood survival hypothesis, assuming an adaptive investment in last eggs to ensure survival of all eggs in the clutch, we found that the size of the last eggs in first clutches was generally larger than the mean egg size of the clutch, and that the relative size of the last egg increased with clutch size. However, a large last egg reflected a general increase in egg size throughout the laying sequence rather than a specific investment in the last egg only. Egg size was not significantly influenced by sex or paternity (within-pair versus extra-pair) of the embryo. In repeat clutches the last egg was not consistently larger than the mean for the clutch. We conclude that female bluethroats face resource limitations during egg formation early in the season, and that the patterns of increase in egg size with laying order for first clutches, and from first to repeat clutches, can largely be explained by proximate constraints on egg formation.     

<Emphasis Type="Italic">Fopius arisanus</Emphasis> oviposition in four <Emphasis Type="Italic">Anastrepha</Emphasis> fruit fly species of economic importance in Mexico

We studied the oviposition performance of Fopius arisanus (Sonan) (Hymenoptera: Braconidae) attacking eggs of four fruit flies of the genus Anastrepha Schiner (Diptera: Tephritidae) under laboratory conditions. The complete process of oviposition on an individual egg of Anastrepha ludens (Loew) lasts in average 85.4 ± 2.9 s, including a tremor (25.8 ± 1.03 s) observed in the middle of this process related to the egg’s descent. The average parasitism of A. ludens egg was 60.9 ± 7.5%, with only 1.2% of superparasitized eggs. During individual acts of oviposition, we noted that F. arisanus possesses a highly flexible ovipositor that curves easily as it searches for additional suitable eggs, which may be of particular benefit when a female finds large clutches of eggs. The individual oviposition of F. arisanus in host fruits attacked by Anastrepha spp. varies with the egg clutch size of each fruit fly species: A. serpentina laid the biggest egg clutches (21.3 ± 1.4), followed by A. ludens (14.2 ± 0.9), and A. striata (1.0 ± 0.0) (=A. obliqua). The time spent by F. arisanus in individual ovipositions was parallel to these findings, reinforcing the idea that F. arisanus attacks several eggs in each individual insertion of its ovipositor. Neither formal oviposition acts, nor adult emergences of F. arisanus were registered in A. obliqua. We discuss the potential of F. arisanus as natural enemy of fruit flies of the genus Anastrepha, and explore the eventual developing of its mass rearing. Handling Editor: Torsten Meiners.     

REPRODUCTION BY NORTHERN BOBWHITES IN WESTERN OKLAHOMA

Abstract: We studied northern bobwhites (Colinus virginianus) in western Oklahoma, USA, during the nesting seasons of 1992–2001. We obtained latitude-specific information on nesting biology and tested hypotheses on the cause of declines in clutch size with progression of the nesting season and on the phenological relation of first, second, and third nesting attempts. For pooled data on bobwhites alive during 15 April-15 September, 64 ± 6.5% of juvenile females (n = 56), 90 ± 10.0% of adult females (n = 9), 13 ± 4.1% of juveniles males (n = 68), and 41 ± 10.7% of adult males (n = 22) incubated ≥ 1 nest. Bobwhites that entered the reproduction period starting on 15 April (n = 229) accumulated 203 nesting attempts (male and female incubations), which translated to 1.7 attempts/hen for all hens that entered (n = 117) and 3.1 attempts/hen for hens that survived to 15 September (n = 65). Overall success for incubated nests (48 ± 2.8%, n = 331) was independent of sex-age class and nesting attempt (1, 2, 3), but it declined at a rate of 2.37%/year (95% CL = 1.10–3.64%/year) during the study. Clutch size declined by 1 egg for every 14–20 elapsed days in the nesting season and the rate of decline was independent of incubation attempt (1 or 2); this result suggests that lower clutch sizes later in the nest season were not necessarily a function of re-nesting. Ending of nest-incubation attempts (1, 2, 3) occurred within an 8-day period from 26 August-2 September. Our results implied that early-season nesting cover is a management concern and that high nest success is possible in the absence of nest predator suppression where abundant nest sites occur across the landscape.     

The breeding biology of Rose-ringed Parakeets Psittacula krameri in England during a period of rapid population expansion

Capsule The reproductive rate of Rose-ringed Parakeets in the UK was higher in 2001-2003 than previously estimated.

Aim To measure reproductive rate and the factors affecting this in Rose-ringed Parakeets in England during a period of rapid expansion in population of this potential invasive species.

Methods During 2001–2003, 108 nests were located and monitored in southwest London, southeast London and the Isle of Thanet. Nest survival using the Mayfield method, fledging success and the characteristics of the nest and surrounding vegetation were measured.

Results The mean date of first egg was 26 March?±?1.3 days and the median clutch size was 4 eggs. Mayfield nest survival rate was 72%. Reproductive success was 1.4?±?0.3 young fledged per nest. Parakeets bred predominantly in ash (Fraxinus) and oak (Quercus). Nest trees had a diameter at breast height (DBH) of 73.7?±?4.7?cm (mean?±?se).

Conclusion Reproductive success in this parakeet was higher than previously estimated (0.8 young fledged per nest prior to 1999). This reproductive output is sufficient to explain the rapid increase in Rose-ringed Parakeet numbers (from an estimated 1500 birds in 1996 to nearly 10,000 birds by 2004) if Rose-ringed Parakeets have a low death rate similar to other parrots.     

Strategies of a successful new invader in European fresh waters: fecundity and reproductive potential of the Ponto-Caspian amphipod Dikerogammarus villosus in the Austrian Danube, compared with the indigenous Gammarus fossarum and G. roeseli

1. Fecundity of a Dikerogammarus villosus population at Spitz was studied in the Austrian Danube during the 3‐year period 2002–2004. Ovigerous females were absent in October and November, and extremely scarce in December when the reproductive season started again slowly. From January to September pre‐copulatory pairs and egg‐carrying females were present. The reproductive cycle lasted for 9–10 months. 2. Various pigmentation phenotypes of D. villosus have been described in the literature. However, no significant differences were found between the reproductive variables studied here and several colour morphs. Mating was size‐assortative; mean body length of males was about 1.3 times greater than that of their potential mates, and the wet weight was approximately twice as heavy. 3. The relationship between the number of embryos per clutch and the wet weight of females was described by a 3‐parameter power equation. The population mean was 43 eggs with a range of five to 194 eggs. Eighty‐two specimens from 1359 D. villosus females had more than 100 eggs: the smallest of these females was 12 mm long (30 mg) wet weight, and the largest, which was 18 mm long (91 mg), had 194 eggs in embryonic development stage 4. 4. Numbers of embryos in developmental stages 2 (early egg stage) and 7 (newly hatched neonates) differed significantly with body wet weight of ovigerous females (P < 0.05). For an average female in the range 10–12 mm (20–30 mg) the number of juveniles in the brood pouch was 74% of the number of stage 2 eggs. This value can be interpreted as the survival rate of eggs. 5. The overall mean egg volume (EV, ±95% CL) of stage 2 eggs of D. villosus was 0.05 ± 0.001 mm³, and EV increased significantly at each stage of development. At stage 6, egg volume had increased by a factor of 2.6, and averaged 0.13 ± 0.001 mm³. In comparison, G. fossarum and G. roeseli had significantly larger eggs in all developmental stages. 6. Mean egg size of D. villosus (0.063 mm³) was maximal in January. For D. villosus (and G. roeseli) the minimum mean egg size occurred in September. In contrast to G. fossarum and G. roeseli, a second peak in egg size was not observed for D. villosus, and egg size fell more or less successively from January to September. 7. A simple index of fecundity was calculated from the number of stage 2 eggs divided by the female's wet weight. The highest values were observed in April and May, when females from the overwintering generation grew to their maximum body size. Thus the release of a large number of neonates corresponds with the availability of plentiful food and rising water temperatures for juvenile growth in the spring. The lowest value occurred in December. In June the small females of a summer generation appeared, with a naturally low fecundity. 8. The relationship between brood development time and water temperature was studied in the laboratory at a series of constant temperatures. At 16 °C, mean brood development time was 14 days for D. villosus, compared with about 3 weeks for the indigenous species. At 10 °C, mean brood development time was 24 days in D. villosus, compared with 40 days in G. fossarum and 44 days in G. roeseli. At 4 °C it was 1.8 and 3.5 times longer in G. fossarum and G. roeseli. 9. The number of offspring produced by a single clutch from a large female D. villosus is considerably higher than the total numbers produced by the indigenous freshwater gammarids, such as G. fossarum, G. roeseli and G. pulex, during their life‐spans of 1.5–2 years in seven to nine successive broods. Only one or two large ovigerous D. villosus would probably be enough to start a new population. A potentially high reproductive capacity, comparatively small eggs, optimal timing to release the maximum number of neonates per female in April/May, and a long reproductive cycle, together with rapid development of eggs, rapid growth to sexual maturation, short life span, tolerance to a wide range of environmental conditions, and exceptional predatory capabilities, all give the invasive Ponto‐Caspian gammarid an opportunity to become globally distributed in freshwater ecosystems of the temperate climate zone.     

Population Irruptions of Northern Bobwhite: Testing an Age-Specific Reproduction Hypothesis

Abstract: Age-specific reproduction has been suggested for northern bobwhite (Colinus virginianus) and has been hypothesized as a factor contributing to population irruptions. However, little research has been conducted on the subject. We conducted a laboratory and field study to determine if age-specific reproduction occurred in northern bobwhites. Our objectives were to compare 7 reproductive measures (% F nesting, date of first incubated nest, egg-laying rate, nesting rate, clutch size, egg mass, and egg hatchability) between first- and second-year breeders and determine if differential reproduction was impacted by diet quality. The laboratory study consisted of a 2 × 2 factorial experiment with age and diet quality (low protein [12%] and high protein [24%]) as the factors. Data for the field study represented a 6-year data set of bobwhite reproduction (May-Sep 2000-2005) obtained from an ongoing radiotelemetry study in southern Texas, USA. We documented similar productivity (i.e., % F laying, egg-laying rate, and egg mass) and timing of laying (i.e., date of first egg) between juvenile (n = 33) and adult bobwhites (n = 27) in our laboratory study. However, females on the high-protein diet exhibited a greater egg-laying rate than females on the low-protein diet. Under field conditions, we also documented no difference in productivity (% F nesting, nesting rate, clutch size, egg hatchability) and timing of nesting (date of first incubated nest) between age classes (n = 59 juv and 32 ad). Our findings do not support early suppositions of age-specific reproduction in quail. Quail irruptions should not be influenced by population age structure as it relates to age-specific reproduction.