Inter-sexual differences in T-cell-mediated immunity of black-headed gull chicks (<Emphasis Type="Italic">Larus ridibundus</Emphasis>) depend on the hatching order

Hatching asynchrony in avian species leads to age and size differences between nestlings within a brood, handicapping last-hatched chicks in the sibling rivalry. Starvation due to this competitive disadvantage has been regarded as the primary cause of an increase in mortality with hatching order. However, for gulls it has also been suggested that disease is the cause of mortality for last-hatched chicks, possibly through reduced immunocompetence and thereby an enhanced susceptibility to infection. In addition, the male-biased mortality reported for several gull species may be related to a higher vulnerability to diseases in males compared to females. To determine the potential influence of the immune system on these mortality patterns, we investigated the T-cell-mediated immunity (CMI) of black-headed gull chicks in relation to hatching order and sex. We found a significant decrease in the CMI with hatching order. This result may be causally related to systematic changes in maternal yolk steroids and carotenoids within the laying sequence. For second-laid eggs, male CMI was significantly lower than female CMI. This is possibly linked to higher plasma levels of testosterone in male embryos which might have an immunosuppressive effect. If so, this effect is masked in eggs of either high (first egg) or low (last egg) quality. Chicks with low CMI showed enhanced mortality rates. Thus the differences in immune response are likely to contribute to the observed mortality patterns. However, hatching order significantly affected mortality independently of CMI, suggesting that competitive disadvantage due to hatching asynchrony is also important.Communicated by M. Webster     

Resource allocation varies with parental sex and brood size in the asynchronously hatching green-rumped parrotlet (<Emphasis Type="Italic">Forpus passerinus</Emphasis>)

When eggs hatch asynchronously, offspring arising from last-hatched eggs often exhibit a competitive disadvantage compared with their older, larger nestmates. Strong sibling competition might result in a pattern of resource allocation favoring larger nestlings, but active food allocation towards smaller offspring may compensate for the negative effects of asynchronous hatching. We examined patterns of resource allocation by green-rumped parrotlet parents to small and large broods under control and food-supplemented conditions. There was no difference between parents and among brood sizes in visit rate or number of feeds delivered, although females spent marginally more time in the nest than males. Both male and female parents preferentially fed offspring that had a higher begging effort than the remainder of the brood. Mean begging levels did not differ between small and large broods, but smaller offspring begged more than their older nestmates in large broods. Male parents fed small offspring less often in both brood sizes. Female parents fed offspring evenly in small broods, while in large broods they fed smaller offspring more frequently, with the exception of the very last hatched individual. These data suggest male parrotlets exhibit a feeding preference for larger offspring—possibly arising from the outcome of sibling competition—but that females practice active food allocation, particularly in larger brood sizes. These differential patterns of resource allocation between the sexes are consistent with other studies of parrots and may reflect some level of female compensation for the limitations imposed on smaller offspring by hatching asynchrony.     

Sex-specific maternal effect on egg mass,laying order,and sibling competition in the Bengalese finch (<Emphasis Type="Italic">Lonchura striata</Emphasis> var. <Emphasis Type="Italic">domestica</Emphasis>)

Maternal effects, such as investment in eggs, have profound effects on offspring fitness. Mothers are expected to skew their investment depending on the laying order and sex when unequal sibling competition occurs within a brood because of sex-specific vulnerability and age hierarchy caused by asynchronous hatching. The Bengalese finch hatches asynchronously and shows a moderate reversed sexual size dimorphism. However, contrary to commonly accepted assumptions of size-dependent vulnerability, the smaller sex (male) is more vulnerable to developmental stress caused by sibling competition. We investigated whether maternal investment would be biased by the position in laying order and the sex of eggs, and also explored the possible differences in growth patterns depending on sex, laying order, and age hierarchy by observing chicks fostered to experimentally manipulated broods where brood composition was controlled and age hierarchy was more enhanced than in natural breeding conditions. We found that overall patterns of maternal investment favored the disadvantageous sectors of sibling competition, i.e., eggs of later laying order and sons over those of early laying order and daughters. We also examined the effect of laying order on adult body size and sex differences in growth patterns. When reared in the subordinate age hierarchy, females could compensate for the deficit of decreased growth rate by taking longer to mature, whereas males could not. We suggest that this sex-specific growth pattern could be the cause of sex differences in vulnerability to early developmental stress.     

Preferential incubation positions for different sized eggs and their influence on incubation period and hatching asynchrony in Snares crested (<Emphasis Type="Italic">Eudyptes robustus</Emphasis>) and yellow-eyed penguins (<Emphasis Type="Italic">Megadyptes antipodes</Emphasis>)

In crested penguins (Eudyptes spp.), second-laid eggs typically hatch before first eggs. Amongst a variety of factors that have been considered as mechanisms underlying this reversal, has been the idea that crested penguins can adjust the degree of hatching asynchrony by manipulating egg positions (i.e. placing the smaller first egg in the supposedly thermally disadvantaged anterior position) during incubation (termed Preferential Incubation Hypothesis). We tested this in the Snares crested penguin (Eudyptes robustus) and the closely related, but synchronously-hatching, yellow-eyed penguin (Megadyptes antipodes). Snares crested penguins were more likely to place their first eggs, which are smaller than second eggs, in the anterior incubation position than were yellow-eyed penguins, which have a clutch of two similar-sized eggs. But when yellow-eyed penguins, a non-brood reducing species, were provided with an artificial size-dimorphic clutch, they also placed smaller eggs more frequently in the anterior position, suggesting that a general preference exists among penguins to place smaller eggs in the anterior position. Egg temperatures of small first eggs of Snares crested penguins were higher in the anterior than in the posterior position. Large first eggs in lesser size-dimorphic clutches experienced high temperature differences in relation to position, while small first eggs in greater size-dimorphic clutches were incubated at similar temperatures. In yellow-eyed penguins, large eggs within clutches generally had higher egg temperatures than small eggs. Incubation periods of second eggs declined with increasing egg size. Egg-size variation, rather than egg positioning behaviour, influenced hatching patterns in Snares crested penguins. In lesser size-dimorphic clutches, second eggs were more likely to hatch first while in greater size-dimorphic clutches, small first eggs were more likely to hatch at the same time or before the second eggs. This was similar in yellow-eyed penguins, where second eggs hatched earlier in clutches with large first eggs. Our data contradicts the Preferential Incubation Hypothesis and we conclude that this hypothesis is unlikely to explain the reversed hatching asynchrony in crested penguins.Communicated by C. Brown     

Cofoundress relatedness and group productivity in colonies of social <Emphasis Type="Italic">Dunatothrips</Emphasis> (Insecta: Thysanoptera) on Australian <Emphasis Type="Italic">Acacia</Emphasis>

Facultative joint colony founding by social insects provides opportunities to analyze the roles of genetic and ecological factors in the evolution of cooperation. Although cooperative nesting is observed in range of social insect taxa, the most detailed studies of this behavior have been conducted with Hymenoptera (ants, bees, and wasps). Here, we show that foundress associations in the haplodiploid social thrips Dunatothrips aneurae (Insecta: Thysanoptera) are most often comprised of close relatives (sisters), though groups with unrelated foundresses are also found. Associations among relatives appear to be facilitated by limited female dispersal, which results in viscous population structure. In addition, we found that per capita productivity declined with increasing group size, sex ratios were female-biased, and some female offspring apparently remained in their natal domicile for some time following eclosion. D. aneurae thus exhibits a suite of similarities with eusocial Hymenoptera, providing evidence for the convergent evolution of associated social and life-history traits in Hymenoptera and Thysanoptera.     

Egg coloration in ring-billed gulls (<Emphasis Type="Italic">Larus delawarensis</Emphasis>): a test of the sexual signaling hypothesis

Although many avian eggs appear to be cryptically colored, many species also lay vibrant blue green eggs. This seemingly conspicuous coloration has puzzled biologists since Wallace, as natural selection should favor reduced egg visibility to minimize predation pressure. The sexual signaling hypothesis posits that blue green egg coloration serves as a signal of female quality and that males exert post-mating sexual selection on this trait by investing more in the nests of females laying more intensely blue green eggs. This hypothesis has received mixed support to date, and most previous studies have been conducted in cavity-nesting species where male evaluation of his partner’s egg coloration, relative to that of other females, may be somewhat limited. In this study, we test the sexual signaling hypothesis in colonially nesting ring-billed gulls (Larus delawarensis) where males have ample opportunity to assess their mate’s egg coloration relative to that of other females. We used correlational data and an experimental manipulation to test four assumptions and predictions of the sexual signaling hypothesis: (1) blue green pigmentation should be limiting to females; (2) extent of blue green egg coloration should relate to female quality; (3) extent of blue green egg coloration should relate to offspring quality; and (4) males should provide more care to clutches with higher blue green chroma. Our data provide little support for these predictions of the sexual signaling hypothesis in ring-billed gulls. In light of this and other empirical data, we encourage future studies to consider additional hypotheses for the evolution of blue green egg coloration.     

Factors affecting vigilance within wild mixed-species troops of saddleback (<Emphasis Type="Italic">Saguinus fuscicollis</Emphasis>) and moustached tamarins (<Emphasis Type="Italic">S. mystax</Emphasis>)

We examine vigilance within a mixed-species troop of saddleback (Saguinus fuscicollis) and moustached (S. mystax) tamarins over a complete year. Saddleback tamarins were consistently more vigilant than moustached tamarins. This may be linked to their preference for lower strata. In accordance with previous studies of other primates, vigilant tamarins of both species were significantly further away from their nearest neighbours, and were also at lower heights in the forest than non-vigilant individuals. There was no observed sex difference in the amount of time spent vigilant. In terms of modes of scanning, the saddleback tamarins looked up significantly more frequently than the moustached tamarins, whereas there was no difference between the species in the frequency of side sweeps. There were no differences between the sexes in the frequencies of either type of vigilant behaviour. The proportion of time spent vigilant was higher than average immediately prior to entering a sleeping site for saddleback tamarins, but not for moustached tamarins. Both species were more vigilant immediately after exiting a sleeping site than at other times of the day. There was significant variation in the amount of time devoted to vigilance over the course of the year. These findings are discussed with respect to the social structure, ecology and main predator threats facing these species.Communicated by D. Watts     

Predation on egg capsules of <Emphasis Type="Italic">Zidona</Emphasis><Emphasis Type="Italic">dufresnei</Emphasis> (Volutidae): ecological implications

Among the diverse patterns of energy allocation to the offspring of gastropods, the presence of egg capsules to protect embryos is common. Females of the edible snail Zidona dufresnei attach egg capsules to hard substrates in shallow Argentine Patagonian waters (40°45′S, 64°56′W) during spring-summer. Embryonic development takes about 30 days at 22°C. In this study, three likely capsule predator species and the marks left by each on egg capsule walls were identified in laboratory experiments in February 2010. Abundances of predators and egg capsules with evidence of predation were assessed in the field in the summers of 2010 and 2011. Under laboratory conditions (N = 10 replicates per treatment and control), the predation rate by the chiton Chaetopleura isabellei was the highest (up to 90%), followed by the gastropod Tegula patagonica and the crab Neohelice granulata (~20% each). Nearly 60% of 41 capsules found in the field showed signs of predation. According to the marks identified in the laboratory, C. isabellei was responsible for 79% of this predation, and T. patagonica for the rest. Predation appears to be important during the encapsulated early life and could be an agent for selecting for resistant capsule walls and a relatively shorter development time.     

Comparative life-history study on sympatric hyperiid amphipods (<Emphasis Type="Italic">Themisto pacifica</Emphasis> and <Emphasis Type="Italic">T</Emphasis>. <Emphasis Type="Italic">japonica</Emphasis>) in the Oyashio region,western North Pacific

Life-history features of the sympatric amphipods Themisto pacifica and T. japonica in the western North Pacific were analyzed based on seasonal field samples collected from July 1996 through July 1998, and data from laboratory rearing experiments. T. pacfica occurred throughout the year, with populations peaking from spring to summer. In contrast, T. japonica were rare from autumn to early winter, but became abundant in late winter to spring. Mature T. pacifica females and juveniles occurred together throughout the year, indicating year-round reproduction. Mature T. japonica females were observed only in spring, and juveniles occurred irregularly in small numbers, suggesting limited, early-spring reproduction in this study area. Size composition analysis of T. pacifica identified a total of eight cohorts over the 2 years of the study. Due to the smaller sample size and rarity of mature females (>9.6 mm) and males (>7.1 mm), cohort analyses of T. japonica were not comparable. Laboratory rearing of specimens at 2°C, 5°C, 8°C and 12°C revealed that a linear equation best expressed body length growth by T. pacifica, while a logistic equation best expressed body length growth by T. japoncia. Combining these laboratory-derived growth patterns with maturity sizes of wild specimens, the minimum and maximum generation times of females at a temperature range of 2–12°C were computed as 32 days (12°C) and 224 days (2°C), respectively, for T. pacifica, and 66 days (12°C) and 358 days (2°C), respectively, for T. japonica. The numbers of eggs or juveniles in females marsupia increased with female body length and ranged from 23 to 64 for T. pacifica and from 152 to 601 for T. japonica. Taking into account the number of mature female instars, lifetime fecundities were estimated as 342 eggs for T. pacifica and 1195 eggs for T. japonica. Possible mechanisms for the coexistence of these two amphipods in the Oyashio region are also discussed.Communicated by O. Kinne, Oldendorf/Luhe     

Between-group encounters among bonnet macaques (<Emphasis Type="Italic">Macaca radiata</Emphasis>)

Socioecological theory suggests that between-group competition is an important factor affecting the nature of primate social relationships. Between-group encounters in macaques may involve female resource defense, male mate defense, and male resource defense. We observed between-group encounters in two groups (a forest group and a temple group) of bonnet macaques (Macaca radiata). We observed 102 encounters in 875 h of observation of the forest group (1.40 per 12-h day) and 58 encounters in 907 h of observation of the temple group (0.77 per 12-h day). Aggressive interactions between groups occurred in 32.4% and 29.3% of encounters in the forest and temple groups, respectively. Overall, we found little support for the female resource defense hypothesis. Females in both groups rarely participated aggressively in between-group encounters. We found support for the male mate defense hypothesis. For example, males of the forest group were more aggressive during encounters in the mating season than in the non-mating season. Males were also aggressive to females from their own group immediately following encounters. We also found partial support for the male resource defense hypothesis. Encounters in the forest group occurred in a feeding context more often than expected based on time budgets. Also, males in the temple group were more often aggressive in food-related encounters than in other encounters. The findings of this study suggest that socioecological models of primate social relationships need to distinguish male and female strategies during between-group encounters and integrate the resulting functional outcomes.Communicated by T. Czeschlik     

Populations of Pecos pupfish (<Emphasis Type="Italic">Cyprinodon pecosensis</Emphasis>) differ in their susceptibility to hybridization with sheepshead minnow (<Emphasis Type="Italic">C. variegatus</Emphasis>)

The importance of sexual selection in hybridization between species that evolved in allopatry is poorly understood. Even less is known about differences between populations in their resistance or susceptibility to genetic introgression with a congener. Here we examine behavioral differences between populations of the native Pecos pupfish (Cyprinodon pecosensis) and their potential effects in facilitating or retarding hybridization with an introduced congener, the sheepshead minnow (C. variegatus). We quantified competitive interactions, establishment of territories, and mating success of several naturally occurring C. pecosensis populations and an artificially created laboratory population (control) against C. variegatus. Experiments were conducted in large outdoor tanks during the summer months when high temperatures favor breeding activity in pupfish. C. pecosensis populations differed in agonistic behavior, territoriality, and in mating success when competing against C. variegatus. Differences in the competitive ability of males from different C. pecosensis populations suggest that the rate of introgression with C. variegatus may vary, depending on the population of C. pecosensis it comes in contact with. Our results suggest that large, stable C. pecosensis populations may be relatively resistant to hybridization with C. variegatus. Interpopulation variation in male behavior, shaped by sexual selection, may be important in determining the incidence and rate of hybridization between the two congeners. Experimental studies are a powerful tool in elucidating behavioral processes, such as the action of intrasexual selection, that facilitate or retard the coalescence of lineages through initial hybridization and subsequent genetic introgression after secondary contact between allopatric species.Communicated by M. Abrahams     

Auditory defence in the peacock butterfly (<Emphasis Type="Italic">Inachis io</Emphasis>) against mice (<Emphasis Type="Italic">Apodemus flavicollis</Emphasis> and <Emphasis Type="Italic">A. sylvaticus</Emphasis>)

Morphological and behavioural traits can serve as anti-predator defence either by reducing detection or recognition risks, or by thwarting initiated attacks. The latter defence is secondary and often involves a ‘startle display’ comprising a sudden release of signals targeting more than one sensory modality. A suggested candidate for employing a multimodal defence is the peacock butterfly, Inachis io, which, by wing-flicking suddenly, produces sonic and ultrasonic sounds and displays four large eyespots when attacked. The eyespots make small birds retreat, but whether the sounds produced thwart predator attacks is largely unknown. Peacocks hibernate as adults in dark wintering sites and employ their secondary defence upon encounter with small rodent predators during this period. In this study, we staged predator–prey encounters in complete darkness in the laboratory between wild mice, Apodemus flavicollis and Apodemus sylvaticus, and peacocks which had their sound production intact or disabled. Results show that mice were more likely to flee from sound-producing butterflies than from butterflies which had their sound production disabled. Our study presents experimental evidence that the peacock butterfly truly employs a multimodal defence with different traits targeting different predator groups; the eyespots target birds and the sound production targets small rodent predators.     

Histological investigation of the reproductive cycles of the limpets <Emphasis Type="Italic">Patella </Emphasis><Emphasis Type="Italic">vulgata</Emphasis> and <Emphasis Type="Italic">Patella ulyssiponensis</Emphasis>

This study devised a staging system for, and monitored, the gonad development of the limpet species Patella vulgata and Patella ulyssiponensis on the South West coast of Ireland using histological techniques. Maturation began in the males of both species in January and in the females it began in March. There was no statistical difference in gonad development between sexes and between species. Spawning in the male P. vulgata occurred from September to December 2003 and in September and October 2004. In female P. vulgata spawning occurred from October to December 2003, no spawning of females was observed in 2004. In male P. ulyssiponensis spawning occurred in November and December 2003 and from September 2004 to December 2004. Spawning was observed from November 2003 to January 2004 and in September 2004 in female P. ulyssiponensis. Sex ratios also varied between the species and between months sampled. Nevertheless more males were observed in both species.     

Genetic analysis of tuna populations,<Emphasis Type="Italic"> Thunnus thynnus thynnus</Emphasis> and<Emphasis Type="Italic"> T</Emphasis>.<Emphasis Type="Italic"> alalunga</Emphasis>

The genetic population structures of Atlantic northern bluefin tuna ( Thunnus thynnus thynnus) and albacore ( T. alalunga) were examined using allozyme analysis. A total of 822 Atlantic northern bluefin tuna from 18 different samples (16 Mediterranean, 1 East Atlantic, 1 West Atlantic) and 188 albacore from 5 samples (4 Mediterranean, 1 East Atlantic) were surveyed for genetic variation in 37 loci. Polymorphism and heterozygosity reveal a moderate level of genetic variability, with only two highly polymorphic loci in both Atlantic northern bluefin tuna ( FH* and SOD- 1*) and albacore ( GPI- 3* and XDH*). The level of population differentiation found for Atlantic northern bluefin tuna and albacore fits the pattern that has generally been observed in tunas, with genetic differences on a broad rather than a more local scale. For Atlantic northern bluefin tuna, no spatial or temporal genetic heterogeneity was observed within the Mediterranean Sea or between the East Atlantic and Mediterranean, indicating the existence of a single genetic grouping on the eastern side of the Atlantic Ocean. Very limited genetic differentiation was found between West Atlantic and East Atlantic/Mediterranean northern bluefin tuna, mainly due to an inversion of SOD- 1* allele frequencies. Regarding albacore, no genetic heterogeneity was observed within the Mediterranean Sea or between Mediterranean and Azores samples, suggesting the existence of a single gene pool in this area.     

Diet of 0-group stages of capelin (<Emphasis Type="Italic">Mallotus villosus</Emphasis>), herring (<Emphasis Type="Italic">Clupea harengus</Emphasis>) and cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) during spring and summer in the Barents Sea

Recruitment of capelin in the Barents Sea fail when juvenile herring and cod are abundant and the potential for feeding competition of wild sympatric capelin and herring larvae and small cod juveniles were investigated. The frequency of gut evacuation after capture of capelin larvae were also studied in mesocosms. Small capelin larvae (<35 mm length) fed on small prey including phytoplankton, invertebrate eggs and nauplii, bivalves, other invertebrate larvae and small copepods. Calanus copepodites were only observed in large capelin larvae (>26 mm length). Calanus copepodites were the major food sources for contemporary herring larvae (25–35 mm length) and Calanus and euphausiids were the major prey for small juvenile herring (37–60 mm length) and cod (18–40 mm length). Capelin larvae reared in mesocosms evacuated the guts shortly after capture. Capelin larvae had a smaller mouth and fed on smaller prey than herring and cod of the same length. This implies that the small capelin larvae, in contrast to sympatric small herring and cod, are not tightly linked to the food chain involving Calanus and euphausiids. Thus, exploitative competition between capelin larvae and planktivorous fish that rely on Calanus and euphausiids in the Barents Sea may be relaxed.     

Disinfection efficacy on cyst viability of<Emphasis Type="Italic"> Artemia franciscana</Emphasis> (Crustacea),<Emphasis Type="Italic"> Hexarthra fennica</Emphasis> (Rotifera) and<Emphasis Type="Italic"> Fabrea salina</Emphasis> (Ciliophora)

Resting stages (cysts) of Artemia franciscana, Hexarthra fennica and Fabrea salina were exposed for two periods (acute and chronic exposures) to different concentrations of five aquaculture disinfectants (formalin, sodium hypochlorite, potassium permanganate, organic iodine, copper sulphate). The effects of chemical treatments were ranked according to their action on cyst hatching: total inhibition (no cyst hatching), heavy inhibition (significant decrease of cyst hatching), alteration of hatching synchrony, and occurrence of anomalous hatching (death of emerging individuals or birth of malformed organisms). Resting-stage viability was not completely impaired by disinfectant exposures at the ordinary doses used in aquaculture, so that usual treatments are possibly ineffective against the protist and metazoan cysts potentially present in breeding systems. On the other hand, the high resistance shown by cysts suggests the possibility of using these chemicals in live-feed disinfection.     

Apparent sibling rivalry in the freshwater fingernail clam<Emphasis Type="Italic"> Sphaerium striatinum</Emphasis>

In Sphaerium striatinum, a freshwater brooding bivalve, up to 97.5% of offspring that adults initially produce fail to reach independence. Marsupial sacs, specialized extensions of gill filaments that act as nurseries, initially contain multiple offspring in various sizes and stages of development. However, by the time offspring reach later stages of development, marsupial sacs typically contain only one offspring. Brood mortality is hypothesized to be the result of competition among embryos for nutrients and/or space. Sphaeriid eggs do not contain enough yolk for offspring to complete development. Adults supply additional nutrients required to reach independence. Brood capacity is limited by adult size. Adults cannot physically brood all offspring they produce. Here, we examine the validity of the competition hypothesis for brood mortality. We reared offspring, in vitro, through metamorphosis under varying nutrient levels and embryo densities. While hatching success and time to hatching were not influenced by nutrients or density, both factors had significant effects on the percentage of embryos completing metamorphosis and timing of metamorphosis. A higher percentage of offspring completed metamorphosis in higher nutrient levels and lower densities. Offspring reared with higher nutrient levels and lower densities also completed metamorphosis more rapidly. We discuss these results in relation to hypotheses for the overproduction of offspring, sibling rivalries, as well as factors that might explain brood mortality in this species.Communicated by T. Czeschlik     

Geographical genetic structure and phylogeography of the <Emphasis Type="Italic">Sargassum horneri</Emphasis>/<Emphasis Type="Italic">filicinum</Emphasis> complex in Japan,based on the mitochondrial <Emphasis Type="Italic">cox3</Emphasis> haplotype

The genetic structure and phylogeography of the brown seaweed Sargassum horneri/filicinum complex in Japan were studied based on the mitochondrial cox3 haplotype. The cox3 haplotypes found were divided into three clades in a statistical parsimony network, among which there were large numbers of steps. Contrary to the reported large amount of drifting S. horneri along the Japanese coast, the three clades were dividedly distributed on the Japanese coast: the northern Pacific, the central Pacific, and western Japan. The western Japan S. horneri had haplotypes that were phylogenetically closer to those of S. filicinum than to the northern and central Pacific S. horneri populations. The S. filicinum populations were included within the western Japan clade and grouped together with the S. horneri samples from western Japan. Taken together with the unstable morphological diagnosis, this result suggests that S. filicinum should be reduced into a synonymy of S. horneri. The TMRCA analysis suggested that the divergence time of each clade may go back to the last interglacial period and a skyline plot suggested that the last glacial maximum had only a small effect on the population size of S. horneri. The geographic subdivision of the three groups, in spite of a large amount of drifting mats, suggests a limited contribution of drifting mats to gene flow on a large geographic scale. On a small geographic scale, a small number of haplotypes were shared between S. horneri-type and S. filicinum-type populations. This result suggests that populations of these two types are partially, though not completely, isolated from each other, possibly by selfing in S. filicinum-type populations or by a difference in peak reproduction.