Hatch or wait? A dilemma in reptilian incubation

Animals often form groups to reduce the risk of predation through the per capita dilution of their individual predation risk. The advantages of grouping also influence the timing of reproduction in many species. In particular, synchrony in the timing of births may have evolved as a predator-avoidance strategy as it dilutes the risk of predation upon vulnerable newborn and naive young. Eggs of an Australian freshwater turtle, Emydura macquarii , can hatch synchronously despite developmental asynchrony among eggs of a clutch and hatchlings have a reduced predation risk by emerging from the nest as a group. Developmental asynchrony within clutches was induced to reflect natural nests by dividing clutches and incubating them at either 25°C or 30°C. Some eggs were then reunited with their clutch-mates and hatching occurred synchronously in some of these groups. In groups where synchronous hatching did not occur, less advanced eggs still hatched earlier than the normal incubation period. Synchrony occurred because the less advanced eggs hatched up to five days earlier than the control embryos. We conclude that the less advanced embryos within a clutch either accelerate their development or hatch prematurely to ensure synchrony of hatching and hatchling group formation may facilitate emergence from the nest and dilute predation risk.     

Age,growth and hatch dates of ingressing larvae and surviving juveniles of Atlantic menhaden Brevoortia tyrannus

Ages, growth and hatch dates of ingressing Brevoortia tyrannus larvae were determined in a 3 year sampling survey at the mouth of the Chesapeake Bay, U.S.A. To determine if otolith‐aged cohorts had variable relative survival, hatch dates of summer‐caught young‐of‐the‐year (YOY) juveniles collected throughout the Chesapeake Bay were compared with hatch dates of ingressing larvae. Modal total length of ingressing larvae was similar among years: 28 mm in 2005–2006 and 2007–2008, and 30 mm in 2006–2007. Ages of ingressing larvae ranged from 9 to 96 days post hatch (dph); mean ages were similar among years, but significantly older in 2006–2007 (50 dph) than in 2005–2006 (44 dph) and 2007–2008 (46 dph). Larval growth rates differed among years. Earliest growth, when larvae were offshore (0–20 dph), was faster in 2006–2007 (0·62 mm day^?1), than in 2005–2006 and 2007–2008 (0·55 mm day^?1 in these years). Subsequently, from 30 to 80 dph, growth was slowest in 2006–2007. Hatch dates of ingressing larvae occurred from September to March and 90% (2007–2008) to 98% (2006–2007) had hatched prior to 31 December. In contrast, most surviving YOY juvenile B. tyrannus had hatched in January to February, suggesting selective mortality of early‐hatched individuals, apparently during the overwinter, larval to juvenile transition period.     

Sex ratio of a population of Anatolian ground squirrelsSpermophilus xanthoprymnus in Central Anatolia, Turkey

We studied the sex ratio of a population of Anatolian ground squirrels,Spermophilus xanthoprymnus (Bennett, 1835), living in a steppe area about 50 km south of Ankara, Central Anatolia, Turkey in 1999–2001. The sex ratio among juveniles and yearlings did not differ from 1∶1, but the sex ratio among adults (≥ 2 years old) was biased toward females. We concluded that from their second summer of life as yearlings onwards, males experienced higher mortality than did females. Such higher mortality is common in ground squirrels, and is likely due to the risks encountered when males disperse from their natal areas.     

CLUTCH SIZE, EGG SIZE, HATCH WEIGHT AND LAYING DATE IN RELATION TO EARLY MORTALITY IN RED GROUSE LAGOPUS LAGOPUS SCOTICUS CHICKS

Clutches of Red Grouse eggs were collected from the wild and subsequent hatching and rearing done in standard conditions in captivity. Variations in chick survival from one clutch to another in the same year were related to differences in hatch weight. Hatch weight was determined only partly by egg size. Weight loss between laying and hatching was related to survival independently of egg size. Variation in this weight loss obscured any simple relationship between egg size and survival, except in eggs laid by captive hens. Intrinsic differences amongst hens caused some variations in laying date, egg size, hatch weight and chick survival. Variations in egg size and hatch weight accounted for less than half the variation in survival; other unmeasured intrinsic factors were also important. Big clutches hatched earlier than small ones. The commonest clutches were of seven and eight eggs, with six and nine frequent. Very big clutches of ten or more eggs were infrequent and chicks from them sometimes survived worse than from smaller clutches. As in other species, the commonest clutch sizes were not the most productive. There was no simple relationship between egg size and clutch size.     

Interannual differences in growth and hatch date distributions of age-0 year walleye pollock Theragra chalcogramma (Pallas) sampled from the Shumagin Islands region of the Gulf of Alaska, 1985 – 2001

Daily increments of age-0 year walleye pollock Theragra chalcogramma otoliths from 1985 to 2001 were examined for interannual variability in growth and hatch dates. Fish were collected in summer and autumn surveys near the Shumagin Islands in the western Gulf of Alaska (GOA). Hatch date distributions of these fish were compared with hatch dates of larvae from spawning aggregations located in Shelikof Strait and the Shumagin Islands based on their spawning times. The hatch date distributions of age-0 year fish captured from the Shumagin Islands area were similar to those observed for larvae from the Shelikof Strait spawning group. Age-0 year fish whose hatch dates corresponded to the Shumagin Islands spawning, which occurred earlier in January and February, were not found. Sea surface temperature was associated with variability in hatch date distribution and growth.     

Preliminary evidence of delayed spawning and suppressed larval growth and condition of the major carps in the Yangtze River below the Three Gorges Dam

Commercial catches of the ‘major carps’ (grass carp Ctenopharyngodon idella, silver carp Hypophthalmichthys molitrix, black carp Mylopharyngodon piceus and bighead carp Aristichthys nobilis) in the middle and lower reaches of the Yangtze River have declined precipitously since construction of the Three Gorges Dam (TGD). To assess specific environmental effects, young-of-the-year major carps were collected below TGD in the Yangtze River at Jianli (larvae) and E’zhou (juveniles) and in the Dongting Lake (juveniles) during June–August 2007. Lapillar otoliths were analyzed to determine early growth. There were no discernible growth increments in otoliths for a majority of the larvae (38 of 63 grass carp and 40 of 47 silver carp), while increments were evident in otoliths formed in the larval stage for all the juveniles, indicating that larvae without discernible increments failed to recruit to the juvenile stage. Back-calculated hatch dates of major carps were between 31 May and 24 July. The initiation of the spawning season was delayed about 1 month compared to pre-TGD records. Larval growth rates, as reflected by otolith-increment width, were lower at Dongting Lake (closer to TGD) than at E’zhou (farther from TGD), indicating that TGD-moderated effects on early growth and development of carps are more apparent for fishes hatched nearer to the dam. The delay in spawning and decreased early growth may partly explain the recent decline of the major carp resources in the middle and lower Yangtze River.     

Selection for synchronous breeding in the European starling

Colonial birds often demonstrate considerable breeding synchrony. In southern Sweden the semi-colonial European starling initiated the vast majority of clutches within one week. Laying dates were positively skewed so that many birds initiated clutches at similar dates early in the season. Breeding was further synchronised by a particularly strong clutch-size reduction equivalent to one third of an egg per day during the first part of the breeding season. The decline in clutch size with season also held true for separate age-classes of females, for individual females laying at different times at different years and for individual females laying at different times the same year. Trends in breeding success during nestling rearing were unlikely to explain the high degree of breeding synchrony or the seasonal decline in clutch size; nestling survival and growth were weakly related or unrelated to reproductive timing. In contrast recruitment success of fledged offspring declined sharply with season. Even within the synchronous laying period, defined as clutches initiated during the first week each year, local recruitment success declined. It is suggested that the early seasonal decline is caused by selection for synchronous fledging permitting the immediate formation of flocks after fledging, whereas the late seasonal trends may be caused by either population differences in female quality or deteriorating conditions for raising young.     

Hatch-period-dependent early growth and survival of Pacific herring Clupea pallasii in Miyako Bay, Japan

Considerable interannual variation in the abundance of larval and juvenile Pacific herring Clupea pallasii was detected in Miyako Bay, on the Pacific coast of northern Japan; abundances were high in 2001 and 2003 and low in 2000 and 2002. Hatch dates and growth rates for larval and juvenile survivors were estimated through otolith analysis. Water temperature and food availability were monitored on the spawning and nursery grounds in the inner part of the bay. The number of spawning females caught in nets set around the spawning ground was recorded during each spawning season (January to May) in 2000–2003. No correlation was found between the number of spawning females and the abundance of larvae and juveniles on the spawning and nursery grounds. The hatch dates of surviving larvae and juveniles were concentrated at the end of the spawning season in 2001 and in the middle of the season in 2003. The larvae experienced relatively high prey concentrations during the first-feeding period in 2001 but low concentrations in 2003. Survival of larvae during the first-feeding period may be a function of prey concentration as well as water temperature. In 2003, low water temperature would reduce starvation mortality during the first-feeding period. In contrast, unfavourable feeding conditions with higher temperatures during the first-feeding period seemed to result in low larval survival in 2000 and 2002. The 2001 larvae grew faster than those in 2003 because of the late hatch dates and the higher ambient temperatures that resulted. Temperature might be a major factor controlling growth rates of C. pallasii larvae in Miyako Bay.     

Temporal dynamics of coral reef fish communities in Nha Trang Bay marine protected area,South-Central Vietnam

To improve understanding of reef fish communities of Vietnam’s first dedicated marine protected area, visual censuses were conducted seasonally from 2003–05 in Nha Trang Bay, south-central Vietnam using SCUBA. Results from this study show that species richness were usually higher in summer than winter and that decreased from summer to winter in both shallow and deep areas in this MPA, but this apparent decrease was not significant. There were significant variations in density of reef fish communities between seasons with higher densities in summer (from April to September) and lower densities in winter (from October to March). The families contributing most to the higher density during summer in the 3 years were acanthurids, chaetodontids, labrids, pomacentrids, scarids, siganids, pomacanthids and caesionids. The increased density of these families in summer was mostly due to increases of juveniles. Recruitment of acanthurids, chaetodontids, labrids, pomacentrids, scarids and siganids occurred primarily in shallow waters whereas caesionids and pomacanthids were mainly recruiting in deeper areas. There were no differences in spatial variations in both species richness and density within the same time periods over several years.     

Male red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>) dispersal during the breeding season

Breeding dispersal can be of significant ecological and evolutionary importance. Yet, it is seldom considered in mammals. I present data on male red deer (Cervus elaphus) movements between sub-populations in southern Sweden during the rut. I investigated whether these movements could be breeding dispersal driven by mate competition. During the ruts of 1998–2009, I recorded 91 movements of males. The longest movement distance was 18.5 km. Dispersal was not restricted to yearlings or sub-adults, but also observed among adult stags. Of 91 movements observed, 7 were made by yearlings, 46 by sub-adults and 38 by adults. There was a significant move among yearlings and sub-adults towards areas with a higher ratio of females/adult males and towards areas with more females. The movements between rutting areas thereby seemed driven by sexual competition.     

Aggression,nesting synchrony,and reproductive fitness in ring-billed gulls

The relationship between synchronous nesting and aggressive behaviour between adults was studied in a colony of ring-billed gulls (Larus delawarensis) from 1976 to 1978. By exchanging eggs and creating more or less synchronous hatching in two of three study plots, I tested the hypothesis that synchronous hatching among neighbouring nests reduces aggression between neighbours. The third plot served as a control. Aggression was lowest in the synchronous plot and highest in the asynchronous plot. More synchronous hatching also reduced aggression when each pair's average synchrony with its neighbours and each neighbour-neighbour interaction were considered. The relationships between neighbour-neighbour hatching synchrony, aggression, and fledging success suggested that gulls that engaged in more aggression had lower reproductive fitness. I discuss the importance of this phenomenon in colonial birds.     

The Number of Louse Eggs on Wild Japanese Macaques (<Emphasis Type="Italic">Macaca fuscata</Emphasis>) Varies with Age,but Not with Sex or Season

During grooming, primates remove harmful ectoparasites, such as ticks and lice, and there is direct evidence for a health benefit of tick removal. Grooming behaviors differ among primates with respect to age and sex. Moreover, the number of ectoparasite may exhibit seasonal variation. Therefore the number of ectoparasites on a host may vary with effects, host age and sex, and season. However, these effects have not been a focus of louse infestation studies of primates. Grooming in Japanese macaques is related to sex and age, with developmental changes in behavior corresponding to the timing of tooth eruption. Moreover, behavioral data for Japanese macaques suggest that lice load may differ with the season. Thus, we examined whether the number of louse eggs varies according to host macaque sex, age, and season, and whether it changes in response to tooth eruption. We counted unhatched and hatched eggs attached to the hair on six 1-cm² areas on the left wrist skin of culled macaques, using a stereoscopic microscope. We sampled five winter coats and three summer coats for each age class: infant, juvenile, adolescent, and adult. The number of unhatched and hatched eggs was related to age, but not to sex and season. There were significant differences in the number of unhatched eggs between infants and adults, juveniles and adults, and adolescents and adults. There were also significant differences in the number of hatched eggs between infants and adults, juveniles and adults, adolescents and adults. Tooth eruption did not influence the number of louse eggs. These results suggest that researchers should consider the age of host animals when assessing the relationship between grooming and ectoparasites.     

Patterns in the abundance,phenology, and hatching of the resting egg stage of the invasive zooplankter <Emphasis Type="Italic">Bythotrephes longimanus</Emphasis>: implications for establishment

To examine how dormancy contributes to the establishment and persistence of Bythotrephes longimanus, we investigated resting egg production and hatching in relation to the demography of the planktonic stage and environmental conditions in Island Lake Reservoir (USA). During a 3-year study, the largest contribution to the egg bank occurred in autumn and most eggs hatched in spring, but we also detected some resting egg production and hatching in summer. The difference between summer and late autumn densities of eggs in sediments averaged 47% (range 0–98%) for 18 sites throughout the reservoir, which was similar to experimental estimates of in situ hatching fraction of 67% for eggs in the spring and summer following their production. Based on emergence traps, neonates hatch in the field during May and June. We estimated mortality rates of 64% for resting eggs and embryos, and 59% for newly emerged neonates. Although hatching fraction saturated at the same level, eggs incubated offshore hatched later than those nearshore where water temperature was warmer and light was detectable at the sediment surface. Low dissolved oxygen concentration did not significantly reduce hatching fraction but resulted in some eggs that initiated development but failed to hatch. Collectively, our results demonstrate substantial annual turnover in the resting egg bank of B. longimanus and high mortality of resting eggs during recruitment from the egg to the first molt of the planktonic stage. These patterns suggest that propagule pressure in the form of resting eggs requires large numbers for establishment, and that considerable post-establishment resting egg production is necessary for inter-annual persistence.     

Synchrony in small mammal populations of montane forest patches in southern India

1. Small mammal populations were studied in montane evergreen forests in the Nilgiris, southern India, using live-trapping from January 1994 to September 1996. Two sites were selected, one with a single large forest patch and adjoining plantations, and the other with several small forest patches separated by grassland.
2. Nine species were recorded, of which eight were trapped in the forest patches, two in the grasslands and five in plantations. Rattus rattus was the most abundant species (2–36 individuals ha⁻¹) followed by Suncus montanus (0–11 individuals ha⁻¹). Densities of seven other species recorded were very low.
3. Synchrony in various population variables – density, biomass, mean weight, density of adults and adult females, and proportion of adults, adult females and sub-adults – was examined for Rattus rattus populations in the forest patches and plantations. Density and biomass were studied in seven other species recorded at these sites. Spearman's rank order correlation was used as a measure of synchrony between the population variables.
4. Within-site synchrony was higher than between-site synchrony in population characters. Synchrony was also higher between plots within the unfragmented site than they were between plots in the fragmented site. Relatively high synchrony in proportions of adults, adult females and juveniles in the forest patches implied that breeding is probably influenced by climate and food availability, which are seasonal in this habitat.
5. Given the small patch sizes (≈1–10 ha) and low population sizes, asynchrony is likely to be an outcome of demographic and environmental stochasticity, and low dispersal rates may impede establishment of synchrony.     

Heterodera glycines Cyst Components and Surface Disinfestants Affect H. glycines Hatching

We investigated the effects of Heterodera glycines cyst components and surface disinfestants on hatching of H. glycines eggs in vitro. Eggs were incubated in either H. glycines cyst wall fragments, cyst wall and egg rinsate, egg homogenate, or control solutions of soybean root diffusate, sterile distilled water, or zinc sulfate. Hatch in cyst wall and egg rinsate, and egg homogenate, was greater (α = 0.05) than hatch in sterile distilled water; however, it was not different from hatch in zinc sulfate according to Dunnett''s test. Hatch in cyst wall fragments was similar to hatch in sterile distilled water. To determine whether surface disinfestants affected hatch, eggs were treated first with chlorhexidine diacetate, mercuric chloride, sodium hypochlorite, or streptomycin sulfate and then incubated in H. glycines egg homogenate, soybean root diffusate, sterile distilled water, or zinc sulfate. Hatch of eggs treated with chlorhexidine diacetate, mercuric chloride, and streptomycin sulfate was reduced (α = 0.05), and hatch of eggs treated with sodium hypochlorite was increased (α = 0.05) relative to hatch of nontreated eggs in all incubation solutions except zinc sulfate according to Dunnett''s Test. Hatch in zinc sulfate was similar among all surface disinfestants except mercuric chloride, where hatch was reduced relative to hatch of nontreated and other surface disinfestant-treated eggs.     

长江口沿岸碎波带刀鲚仔稚鱼的日龄组成与生长

实验研究了长江口沿岸碎波带刀鲚仔稚鱼的日龄组成、孵化期、早期生长率和滞留时间。从2007年5-10月在长江口沿岸碎波带采集的刀鲚仔稚鱼中,共选取594尾(体长范围为3.0-30.5mm),划分发育阶段,并取矢耳石进行日龄分析。仔稚鱼日龄范围为7-34d,以13-18d日龄比例较高,占总数的50.1%;仔稚鱼体长(L,mm)与日龄(D,days)呈显著直线关系:L=0.73D+5.09,R2=0.74;孵化期为5月23日至10月4日,高峰期集中在5月末至8月上旬,且早期个体在孵化后7d左右开始进入到碎波带,在碎波带滞留约23d。       

Differences in egg size, shell thickness, pore density, pore diameter and water vapour conductance between first and second eggs of Snares Penguins Eudyptes robustus and their influence on hatching asynchrony

Brood reduction in birds is frequently induced by hatching asynchrony. Crested penguins (genus Eudyptes) are obligate brood reducers, but in contrast to most other birds, first‐laid eggs are considerably smaller in size than second‐laid eggs; furthermore, first‐laid eggs hatch after their siblings. The mechanisms underlying this reversal in size and hatching order remain unclear. In this study, we tested whether the second‐laid eggs of Snares Penguins Eudyptes robustus have a higher eggshell porosity allowing them to maintain a higher metabolic rate throughout incubation and to hatch before their first‐laid siblings. We investigated differences in egg size, shell thickness, pore density, pore diameter and water vapour conductance between first and second eggs within clutches and examined the influence of these shell characteristics on hatching asynchrony. First‐laid eggs of Snares Penguins were approximately 78% of the size of the larger second eggs. Second‐laid eggs had considerably thicker shells and more pores per cm² than first eggs, whereas pore diameter did not differ between eggs. Water vapour conductance was greater in second‐ (16.8 mg/day/torr) than in first‐laid eggs (14.9 mg/day/torr). The difference in water vapour conductance between first‐ and second‐laid eggs within clutches was related to hatching patterns. In nests where second eggs hatched before first‐laid eggs, second eggs had a considerably greater water conductance than their sibling, whereas in nests where both eggs hatched on the same day, the difference in water conductance between eggs was very small, and in a few nests where small first eggs hatched before their larger sibling, they had a greater water conductance than their larger second‐laid nestmate. Surprisingly few studies have investigated differences in shell characteristics between eggs within clutches and associated effects on hatching asynchrony. This study has demonstrated that such differences exist between eggs within clutches and that they can influence hatching patterns.     

Spatial dynamics in breeding performance of a predator: the connection to prey availability

Using nationwide long-term data on goshawk and grouse populations in Finland we study the spatial dynamics of the numbers of breeding northern goshawk ( Accipiter gentilis ) pairs, goshawk brood size and offspring sex ratio and their connection to the abundance of grouse. Our first large-scale data comprise of observations on goshawk nests during 1986–2001 pooled to 21 different regions. The second set are annual (1989–1998) observations of brood size and offspring sex ratio (females over the sum of females and males) in goshawk nests all over the country, aggregated to 50 km grid level (n=28 grid units). The third set comprises counts (1989–2001) of four species of woodland grouse, split to adults and juveniles, also given in the same 50 km grid units. Using these data, we show that the annual numbers of northern goshawk nests in the different regions fluctuate in synchrony. Synchrony is also found in long-term fluctuations of northern goshawk brood size and offspring sex ratio. Moreover, synchrony is found in annual numbers of grouse juveniles and adults, the main prey for the northern goshawk. In the brood size and offspring sex ratio of the goshawk, as well as in the annual numbers of grouse juveniles and adults the degree of synchrony falls off with increasing distance. However, only in sex ratios and in grouse dynamics are the slopes of synchrony vs distance roughly matching. We also found that sex ratio either vs grouse juveniles or grouse adults has a more matching spatial dimension (50 km radius) that sex ratio vs brood size. These observation lend support to the hypothesis that goshawk offspring sex ratio and grouse abundance are interconnected. Despite the reason, consequences of spatial coupling in sex ratio could have repercussions on other life history events.     

Fitness Components and Selection of Biotypes of Heterodera glycines

Survival of biotypes of Heterodera glycines was studied in microplots and in the field. The field population was subjected to various cropping sequences. Viability of eggs overwintered in microplots was determined each spring by percentage hatch, percentage of hatched eggs penetrating roots, and numbers of females developing on Peking and PI 88788 soybeans. Eggs from the field were collected in the spring and fall and assayed for ability to develop on Peking and PI 88788. Hatch of isolates overwintered in the microplots averaged 13% in May 1989 and 19% in 1990. No differences in hatch were detected among the isolates in 1989. Numbers of juveniles penetrating susceptible roots averaged less than 20% of the hatched eggs each year. An isolate of a biotype parasitic on susceptible soybeans and the resistant soybean PI 88788 penetrated roots more successfully than other biotypes. A second isolate from North Carolina, parasitic on susceptible soybeans, PI 88788, and the resistant soybean Peking experienced selection against development on Peking during two winters. Only 17 % of the expected numbers of females developed on Peking from this isolate. In the microplot experiment, parasitism of PI 88788 and Peking had a selective disadvantage (selection coefficient) of s = 0.29 and 0.62 over all isolates, respectively. In the field experiment, the relative numbers of cysts on Peking and PI 88788 increased between the spring and fall on soybean, then decreased over the winter and under corn. Selection coefficients against parasitism of PI 88788 and Peking averaged 0,19 and 0.3 in the field population. In neither experiment did juveniles lose their ability to parasitize susceptible soybeans.