Reinforcement in the banded darter Etheostoma zonale: The effect of sex and sympatry on preferences

Reinforcement occurs when selection against hybrid offspring strengthens behavioral isolation between parental species and may be an important factor in speciation. Theoretical models and experimental evidence indicate that both female and male preferences can be strengthened upon secondary contact via reinforcement. However, the question remains whether this process is more likely to affect the preferences of one sex or the other. Males of polygynous species are often predicted to exhibit weaker preferences than females, potentially limiting the ability for reinforcement to shape male preferences. Yet, in darters (Percidae: Etheostoma), male preference for conspecific mates appears to arise before female preferences during the early stages of allopatric speciation, and research suggests that male, but not female, preferences become reinforced upon secondary contact. In the current study, we aimed to determine whether the geographically widespread darter species Etheostoma zonale exhibits a signature of reinforcement, by comparing the strength of preference for conspecific mates between populations that are sympatric and allopatric with respect to a close congener, E. barrenense. We examined the strength of preference for conspecifics for males and females separately to determine whether the preferences of one or both sexes have been strengthened by reinforcement. Our results show that both sexes of E. zonale from sympatric populations exhibit stronger conspecific preferences than E. zonale from allopatric populations, but that female preferences appear to be more strongly reinforced than male preferences. Results therefore suggest that reinforcement of female preferences may promote behavioral isolation upon secondary contact, even in a genus that is characterized by pervasive male mate choice.     

Early development of the rainbow darter,Etheostoma caeruleum,according to the theory of saltatory ontogeny

Synopsis The early development of rainbow darter, Etheostoma caeruleum, was examined from an ecological perspective. Steps and thresholds of ontogeny to completion of body squammation are defined, and related to environmental factors. Rainbow darter eggs are about 2 mm diameter, considerably larger than those of related logperch (Percina caprodes). The embryonic vitelline respiratory plexus is much more extensive. The pelagic interval characteristic of logperch and ancestral percids is eliminated and onset of exogenous feeding is delayed. The larger larvae of the rainbow darter can begin feeding directly on aquatic insects, and complete their life cycle in streams. Therefore, shifts in the timing of important thresholds (e.g. exogenous feeding) are ecologically important. Furthermore, early maturation and/or delayed bone and scale formation may be responsible for reductions in the lateralis system and scalation in this and other darter species.     

Early development of johnny darter,Etheostoma nigrum,and fantail darter,E. flabellare,with a discussion of its ecological and evolutionary aspects

Synopsis Fantail darters,Etheostoma flabellare, have larger eggs with greater yolk volume than johnny darters,E. nigrum, and have a longer embryo period. This longer embryo period is associated with accelerated finfold differentiation, producing a larger, better formed individual at the onset of exogenous feeding. Developmental differences among these two species and the rainbow darter,Etheostoma caeruleum, primarily in timing of ontogenetic events, are described in detail. These differences in timing are considered largely early developmental phenomena as they affect embryos, larvae, and aievins but apparently not juveniles or adults. Demographic aspects of early life history were assessed using data from studies of Ontario streams. Differences in early survival among species appear to play an important role in recruitment of mature females, but cannot be attributed to parental care or egg size differences.     

PERMANENT GENETIC RESOURCES: Microsatellite DNA primers for the candy darter, Etheostoma osburni and variegate darter, Etheostoma variatum, and cross-species amplification in other darters (Percidae)

In order to investigate a potential hybrid zone between the candy darter, Etheostoma osburni, and variegate darter, Etheostoma variatum, and examine population variation within E. osburni, a suite of primers for 15 polymorphic microsatellite loci were developed. The average number of alleles per locus was 5.5 in E. osburni and 7.6 in E. variatum, and the average observed heterozygosities were 62.5% and 71.4%, respectively. There were no deviations from Hardy-Weinberg equilibrium and no observed linkage disequilibrium after Bonferroni correction. The utility of these primers was also tested in 11 species of darters representing all four genera of darters. Success of cross-species amplification was largely consistent with phylogenetic relationships of darters.     

Annual variation in the seasonal shift in egg size and clutch size in Sceloporus woodi

Summary Early and late season clutch parameters were examined over a three year period in the Florida scrub lizard, Sceloporus woodi. Precipitation levels were monitored throughout the study. In the early and late season of 1984 and the early season of 1986 precipitation levels approximated long-term mean levels of precipitation. In 1985 a severe winter drought occurred. Clutch size was positively related to body size in all samples in every year. In 1984 and 1986, egg size was not related to clutch size, whereas, in 1985 egg size was negatively related to clutch size. In 1985, females produced large clutches of small eggs early in the season and small clutches of large eggs late in the season. In 1984, no seasonal changes in egg or clutch size occurred. In the late season of 1986, females produced the largest clutches and the smallest eggs of all the samples, but egg and clutch size were not statistically different from the early season egg and clutch size of 1986. Total clutch dry weight, an estimate of total clutch energy, was not different in any of the six sampling periods. These data do not support current adaptationist models that attempt to explain the control of clutch and egg size in lizards. It is argued in this paper that egg and clutch size may vary in response to past environments that affect a female's physical condition, as well as, current resources that may be important for maintenance and reproduction. Egg and clutch size appear to be plastic traits selected to respond to proximal environmental variation, whereas, the investment of total dry matter/clutch has been optimized.     

Evidence of divergent growth rates among populations of the lizard Anolis carolinensis based on experimental manipulations of egg size

Geographic variation in body size is of special interest because it affects nearly all aspects of an organism’s life. I examined whether differences in body size among four populations of the green anole lizard, Anolis carolinensis, were attributable to maternal investment in egg size and/or growth rates of embryos and juveniles. Larger body size and larger egg size relative to female size in the northern part of the range have been documented in this species, and suggested to be adaptive responses to more extreme winters. The current study confirmed the trends in adult size and egg size in the north, but rejected the trend of larger egg size relative to body size in the south. To control for differences in maternal investment in egg size among populations, I performed yolk removals on eggs from two northern populations to produce comparably sized eggs relative to one southern population. This manipulation was designed to minimize the confounding effect of maternal investment in yolk, the primary energy reserves for eggs, so that intrinsic differences in embryonic growth due to metabolism could be investigated. I found that differences in juvenile and, potentially, embryonic growth rates existed among populations of A. carolinensis, both due to and independent of differences in egg size. Juveniles from the northernmost population were bigger not only due to larger egg size, but also due to faster juvenile growth and possibly differences in developmental stage of oviposition or conversion of egg mass to hatchling mass. Larger body size may hold a number of advantages in northern populations of this species, including starvation resistance through winters and better competitive access to food resources and warmer microhabitats.     

Isolation and identification of eight microsatellite loci in the Cherokee darter (Etheostoma scotti) and their variability in other members of the genera Etheostoma, Ammocrypta, and Percina

The Cherokee darter Etheostoma scotti is a federally threatened fish endemic to the Etowah River system of northwest Georgia. In order to analyse the population structure and genetic diversity of this fish, eight tetranucleotide microsatellite genetic markers were developed. The marker set was applied to 13 additional darter species to test cross-species amplification and polymorphism. Successful amplification was obtained for all eight loci in each of the 13 other species of darters, with between seven and eight polymorphic loci per species.     

Characterization of eight polymorphic microsatellite DNA markers for the greenside darter,Etheostoma blennioides (Percidae)

The greenside darter, Etheostoma blennioides is a small benthic fish found in fast‐flowing streams in eastern North America. In Canada, this species is native to three, and introduced into one, Great Lakes tributaries in southwestern Ontario. It is currently listed as a species of Special Concern. To characterize population genetic structure and diversity in the Canadian populations of greenside darter, eight polymorphic microsatellite markers were developed for the species. The polymerase chain reaction primers were tested between 32 and 60 individuals from the Sydenham River and yielded a high number of alleles (four to 42 per locus), and observed heterozygosities ranging from 0.14 to 0.82.     

Reproductive biology and spawning habitat supplementation of the relict darter, Etheostoma chienense, a federally endangered species

We investigated the reproductive biology of Etheostoma chienense in the Bayou du Chien drainage of western Kentucky. Etheostoma chienense is similar in its ecology and reproductive biology to other members of the E. squamiceps complex. However, E. chienense is opportunistic in its choice of egg deposition substrates, as nests were found on a variety of natural and anthropogenic items. Due to an apparent lack of suitable spawning substrates, we added half-cylindrical ceramic tiles to several stretches of stream to increase potential nest productivity. Egg-clutches attached to artificial substrates were twice the size of egg clutches attached to naturally occurring materials. Laboratory experiments were conducted to determine nest rock size and mate choice preferences. Both the size of the male and of the nest rock appeared to be important parameters to spawning females. The life history information gathered here has significant management implications. Artificial spawning substrates should be placed in appropriate microhabitats (i.e., shallow, low-flow reaches in headwaters). Nest cavity vertical height should be about 3.0 cm and tiles should be spaced at least 0.5 m apart. We feel the use of surrogates (e.g., E. oophlyax) to investigate other types of spawning cover, and restoration of riparian buffer zones among other actions, would particularly benefit recovery efforts for this endangered species.     

Clutch production in the darter Etheostoma lynceum and its implications for life-history study

We tested two hypotheses that describe previously untested assumptions about the appropriate stage(s) for use in determining clutch sizes in darters. Three sequential stages of clutch/ovarian development are recognized: mature oocyte/ovary, ripening oocytc/ovary, and ripe oocyte/ovary. Mean ratios of clutch size to female length (relative clutch size, RCS) were significantly smaller and variances of RCS were significantly greater in ripe females when compared to mature and ripening females. Correlation coefficients for the relationship between clutch size and standard length (S.L.) were significantly greater for mature and ripening females than for ripe females. Mean clutch size, adjusted for the S.L. covariate, was significantly greater in mature and ripening females than in ripe females. Thus, we conclude that counts of eggs from ripe females yield fecundity estimates that are lower and more variable than estimates from counts of oocytes from mature and ripening females collected at the same time. Based on this conclusion, we discuss methods for developing fecundity estimates in darters. Our results may be broadly applicable to other taxa of fishes with group-synchronous ovum development.     

Spawning and nesting behavior of the waccamaw darter,Etheostoma perlongum

Synopsis The spawning and nesting behavior ofEtheostoma (Boleosoma) perlongum was investigated in the field and laboratory. Sexual dimorphism is highly developed in such features as genital papillae, first dorsal and paired fins, and nuptial coloration. A reproductive migration from mid-lake to shore occurs in the spring: males precede females to select nest sites under submerged sticks and other debris. The male excavates a depression beneath the submerged object. Gonad analysis indicates a single spawning season extending from March through June. Nests were found from late April to mid-June and were guarded by a single male for periods of 13 to 36 days. Males initiate courtship by lateral display, lead the female to the nest site and show the nest by inverting. The female responds by tail up, tail wag and circle; males also tail wag and circle. Spawning pairs invert, usually in unison, and orient head to head or, less often, head to tail. The female deposits eggs while holding her body in a weak S or J shape with the caudal peduncle held away from the spawning substrate while vibrating.     

Clutch size variation in the Nazca booby: a test of the egg quality hypothesis

In obligately siblicidal bird species, aggressive behavior bya dominant chick results in a fixed brood size of one, yetthese species usually show clutch size variation between individuals.Simmons proposed that variation in clutch size in obligatelysiblicidal species is related to a trade-off between egg qualityand egg quantity: some individuals produce a single highly hatchable egg, while others produce two small, lower qualityeggs. We tested the egg quality hypothesis as an explanationfor observed clutch size variation in the Nazca booby (Sulagranti), an obligately siblicidal seabird. We tested the assumptionthat egg volume is positively correlated with hatchabilityand the prediction that eggs from one-egg clutches are largerthan eggs from two-egg clutches. We did not find a positive relationship between egg volume and hatchability in this species.Eggs from two-egg clutches were either equivalent in volumeor larger than eggs from one-egg clutches. Thus, the egg qualityhypothesis was rejected as an explanation for clutch size variationin the Nazca booby. Instead, two-egg clutches appear to befavored because of the insurance value of the second-laid egg,while one-egg clutches result from food limitation.     

Life history of the gulf darter,Etheostoma swaini (Pisces: Percidae)

Synopsis Etheostoma swaini, the gulf darter, was collected from the Black Creek drainage in southern Mississippi (February 1978 – April 1979). The gulf darter generally inhabits small- to moderate-size creeks and occurs over a sand or sandy mud bottom, often in association with aquatic vegetation or a layer of organic debris. Larval dipterans were the most important food items, both numerically and volumetrically. Chironomids were found in 71–100% of the stomachs in all except the unusual March 16 collection. The length frequency distribution and the scale annuli analysis indicated there were three year-classes present in the population at any one time. Fifty-one percent of the specimens taken were less than 12 months old. During the mid-February to late March spawning season gulf darters were most often collected over clean gravel or gravelsand substrates. Laboratory observations suggest that the female burrows into the gravel where the demersal, adhesive eggs are deposited. Female gulf darters significantly outnumbered males at a ratio of 59:41.     

Clutch size, intraclutch egg size variation, and breeding strategy in the GoldcrestRegulus regulus

Summary In Fennoscandia the Goldcrest regularly lays two clutches per breeding season. Near Trondheim in Central Norway (about 63° N) the mean size of 11 first clutches was 10.4 (9–12) eggs and of 10 second/repeat clutches 9.4 (8–11) eggs. Mean egg length was 13.49 mm, breadth 10.35 mm, mean weight of freshly-laid eggs 0.78 g, volume 737 mm³ and shape index 130.6. The correlation coefficient for egg length against weight was 0.47, and for egg breadth against weight 0.77. For the four clutches studied in more detail, all egg dimensions, except the shape index, increased with the laying sequence. On average, the last-laid egg was 20.1 % heavier than the first-laid egg, or 5.6 % heavier than the mean weight for the whole clutch. The 3–5 last-laid eggs in the clutch deviated by about the same magnitude from the clutch mean. Because of the pronounced asynchrony in hatching, the size-hierarchy within a Goldcrest brood may be considerable, at least 1:5. The survival chances of the smallest young in a brood apparently depend mainly on the prevailing food supply. The primary advantage of hatching from a relatively large-sized egg is probably that such a hatchling is strong enough to secure itself a favourable feeding position in the crowded nest, in which the young often form 2–3 horizontal layers. The parental selection of food is adjusted to the size of the young, even in a brood with a marked size-hierarchy. It is concluded that the Goldcrest has evolved a brood reduction strategy.

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Gelegegröße, Variation der Eigröße und Brutstrategie beim Wintergoldhähnchen (Regulus regulus)

Zusammenfassung In Fennoskandien zeitigt das Wintergoldhähnchen regelmäßig zwei Gelege pro Brutsaison. In einem Kontrollgebiet bei Trondheim in Mittelnorwegen (ca. 63° N) betrug die mittlere Gelegegröße für 11 Erstgelege 10,4 (9–12) und für 10 Zweit- bzw. Ersatzgelege 9,4 (8–11) Eier. Die mittlere Eilänge betrug 13,5 mm, die Eibreite 10,35 mm, das mittlere Frischvollgewicht 0,78 g; das Volumen maß 737 mm³, der Eiformindex (=Länge/Breite × 100) 130,6. Die Korrelation zwischen Länge und Gewicht betrug 0,47, zwischen Breite und Gewicht 0,77. Bei vier genauer untersuchten Gelegen nahmen alle Eidimensionen mit Ausnahme des Eiformindex mit der Legefolge zu. Im Mittel war das letzte Ei 20,1 % schwerer als das erste oder 5,6 % schwerer als das mittlere Gewicht des gesamten Geleges; letzteres gilt auch für die 3 bis 5 zuletzt gelegten Eier. Als Folge des bemerkenswert asynchronen Schlupfes sind die Größenunterschiede innerhalb einer Goldhähnchenbrut beachtlich, nämlich mindestens 1:5. Die Überlebenschancen des kleinsten Jungen einer Brut hängen offensichtlich vor allem vom Nahrungsangebot ab. Der primäre Vorteil eines Jungen aus einem großen Ei liegt möglicherweise darin, daß dieser Nestling stark genug ist, um sich eine günstige Position im dichtbesetzten Nest bei den Fütterungen zu sichern. Die Jungen liegen oft in zwei bis drei Schichten übereinander. Die Nahrungswahl der Altvögel richtet sich nach der Jungengröße, sogar in einer Brut mit deutlichen Größenunterschieden. Das Wintergoldhähnchen hat wahrscheinlich die Strategie einer Brutreduzierung in Anpassung an knappes Nahrungsangebot entwickelt.

     

Intraspecific egg size variation and sperm limitation in the broadcast spawning bivalve Macoma balthica

Bottom-ice algae within Antarctic sea ice were examined using chlorophyll fluorescence imaging. The detailed structure of the bottom-ice algal community growing in the platelet and congelation layers of solid pieces of sea ice was evident for the first time in chlorophyll imaging mode. Strands of fluorescence representing algal cells were clearly visible growing upward into brine channels in a fine network. Images of effective quantum yield (Ф_PSII) revealed that the Ф_PSII of algae embedded in the sea ice was approximately 0.5. Furthermore, Ф_PSII decreased slightly with distance from the ice-water interface.The response of Antarctic sea ice algae to changes in irradiance and salinity, and the effects of slowly warming and melting the ice block sample were examined using this system. The Ф_PSII of bottom-ice algae decreased as irradiance increased and salinities decreased. Bottom-ice algae appear to be most vulnerable to changes in their environment during the melting process of the ice, and this suggests that algae from this region of the ice may not be able to cope with the stress of melting during summer.Chlorophyll fluorescence imaging provides unprecedented imagery of chlorophyll distribution in sea ice and allows measurement of the responses of sea ice algae to environmental stresses with minimal disruption to their physical habitat. The results obtained with this method are comparable to those obtained with algae that have been melted into liquid culture and this indicates that previous melting protocols reveal meaningful data. In this chlorophyll imaging study, rapid light curves did not saturate and this may prevent further use of this configuration.     

Offspring fitness in relation to population size and genetic variation in the rare perennial plant species Gentiana pneumonanthe (Gentianaceae)

Seeds were sampled from 19 populations of the rare Gentiana pneumonanthe, ranging in size from 5 to more than 50,000 flowering plants. An analysis was made of variation in a number of life-history characters in relation to population size and offspring heterozygosity (based on seven polymorphic isozyme loci). Life-his-tory characters included seed weight, germination rate, proportion of seeds germinating, seedling mortality, seedling weight, adult weight, flower production per plant and proportion of plants flowering per family. Principal component analysis (PCA) reduced the dataset to three main fitness components. The first component was highly correlated with adult weight and flowering performance, the second with germination performance and the third component with seed and seedling weight and seedling mortality. The latter two components were considered as being maternally influenced, since these comprised life-history traits that were significantly correlated with seed weight. Multiple regression analysis showed that variation in the first fitness component was mainly associated with heterozygosity and not with population size, while the third fitness component was only correlated with population size and not with heterozygosity. The latter relationship appeared to be non-linear, which suggests a stronger loss of fitness in the smallest populations. The second (germination) component was neither correlated with population size nor with genetic variation. There was only a weak association between population size, heterozygosity and the population coefficients of variation for each life history character. Most correlation coefficients were negative, however, which suggests that there is more variation among progeny from smaller populations. We conclude that progeny from small populations of Gentiana pneumonanthe show reduced fitness and may be phenotypically more variable. One of the possible causes of the loss of fitness is a combination of unfavourable environmental circumstances for maternal plants in small populations and increased inbreeding. The higher phenotypic variation in small populations may also be a result of inbreeding, which can lead to deviation of individuals from the average phenotype through a loss of developmental stability.     

Clutch size in parasitoids: the egg production rate as a constraint

Summary In many species of insect parasitoids, adult females mature eggs as they search their environment for hosts. In such species, the number of mature eggs, at the point of finding a host, is a function of the interhost time and the rate of egg maturation. Assuming that interhost search times are variable, we use a version of the marginal value theorem to derive a decision rule for optimizing the time spent exploiting individual hosts; this indirectly determines clutch size. We find that a threshold search time exists above which a female should simply lay her currently mature eggs and depart from the host. However, when the search time has been less than the threshold, a female should oviposit, but then remain on the host to mature and lay additional eggs, until the threshold time is reached.     

Clutch and egg size in ladybird beetles

There was a significant positive correlation between the number of ovarioles and clutch size in the 8 species of cocinellid studied. There was no intraspecific correlation between egg size and clutch size under standard conditions. These tactics are discussed in terms of the allocation of resources to reproduction.      

Geographic variations in life-history traits of the intertidal ocypodid crab Macrophthalmus banzai

Reproductive traits of the mud crab Macrophthalmus banzai were compared in populations from Okinawa (26°13′ N) and Amakusa (32°32′ N). Crabs matured earlier and at a smaller size in Okinawa than in Amakusa, but growth rate did not differ between the two localities. Population density was much higher in Okinawa. Breeding occurred in winter in Okinawa and in summer in Amakusa. In Okinawa, females produced one brood at over 1 year of age and then died. In Amakusa, females produced one brood at over 1 year of age, and one or two broods at around 2 years of age, and then died. Egg weight in the Okinawa population was almost twice that of the Amakusa population. Furthermore, brood weight was larger in Okinawa than in Amakusa. Longevity was 16–19 months in Okinawa and 28–30 months in Amakusa. At high latitudes, the winter low temperature blocks reproduction of M. banzai. At low temperature blocks reproduction of M. banzai. At low latitudes, their short lifespan prevents iterative breeding. Thus, the difference in breeding season greatly affects other life-history traits.     

A resource budget model to explain intraspecific variation in mast reproductive dynamics

Rowan, Sorbus aucuparia L., exhibits conspicuous geographic variation in its inter-annual patterns of reproduction across southern Norway. Along the southern west coast, trees flower and produce clusters of succulent berries every other year, while in the east, trees mast every 3 years. We investigate two hypotheses for this variation: local adaptation to environmental conditions that favors different reproductive schedules at different locations or variation arising from single stereotyped reproductive strategy in the face of a geographic variation in productivity. To assess the theoretical plausibility of each of these hypotheses, we develop a refined resource budget model for mast reproduction. The refined model assumes that there is a lower bound on the amount of resource that may be allocated to reproduction, which allows the prediction of the relation between reproductive schedules and productivity. From the analysis of the model, we find that the observed geographic transition in rowan mast is caused if either productivity in the east is lower than that in the west or if plants in the eastern populations have evolved to pause reproduction until energy reserves are replenished to higher levels. The prediction by the productivity gradient hypothesis match with the empirical finding that the productivity is, in fact, likely to be lower in the east than the west, although we lack empirical data to test the likelihood of the local adaptation hypothesis. There is a need for experimental studies to clarify the validity of the local adaptation hypothesis. Akiko Satake is the recipient of the 11th Denzaburo Miyadi Award.