Book Review

Abstract

Most studies on population ecology of bryophytes have involved common species. However, some studies have compared life history strategies in rare and common species. We review the life history strategies (life expectancy, sexual and asexual reproduction, spore production, spore size and dispersal) for species that are rare in relation to pattern and persistency of suitable habitat patches. In particular, we discuss the dynamics on two levels, within and among localities, for different categories of rare species. We predict that most rare species will be found to have restricted dispersal capacities but higher than average life expectancies of local subpopulations. Natural rarity is distinguished from human-induced rarity and species rare for the latter reason are distinguished as 'threatened'.     

Life‐history syndromes: Integrating dispersal through space and time

Recent research has highlighted interdependencies between dispersal and other life‐history traits, i.e. dispersal syndromes, thereby revealing constraints on the evolution of dispersal and opportunities for improved ability to predict dispersal by considering suites of dispersal‐related traits. This review adds to the growing list of life‐history traits linked to spatial dispersal by emphasising the interdependence between dispersal through space and time, i.e. life‐history diversity that distributes individuals into separate reproductive events. We reviewed the literature that has simultaneously investigated spatial and temporal dispersal to examine the prediction that traits of these two dispersal strategies are negatively correlated. Our results suggest that negative covariation is widely anticipated from theory. Empirical studies often reported evidence of weak negative covariation, although more complicated patterns were also evident, including across levels of biological organisation. Existing literature has largely focused on plants with dormancy capability, one or two phases of the dispersal process (emigration and/or transfer) and a single level of biological organisation (theory: individual; empirical: species). We highlight patterns of covariation across levels of organisation and conclude with a discussion of the consequences of dispersal through space and time and future research areas that should improve our understanding of dispersal‐related life‐history syndromes.     

The trade-off between vegetative and generative reproduction among angiosperms influences regional hydrochorous propagule pressure

Aim Local communities are subject to spatiotemporal contingencies of landscape processes; community assembly is thus often considered to be unpredictable and idiosyncratic. However, evolved trade‐offs of species’ life histories may set distinct constraints on the assembly of species communities. In plants, the recruitment and invasion success of species into communities depend primarily on the number of propagules available and on their generative or vegetative character. Life‐history trade‐offs prevent individual plants from producing large numbers of both generative and vegetative propagules, but it is not clear whether this constrains their availability at the landscape scale. We thus tested whether: (1) the observed relationship between generative and vegetative propagules deviates from the null expectation stating that species contributing the bulk of generative propagules to the propagule rain should also contribute the bulk of vegetative propagules; and (2) whether vegetative and generative propagule pressures are negatively correlated once species abundance in the regional pool is accounted for. Location A large riparian landscape in the Netherlands. Methods Analyses were based on an extensive trapping of floating propagules (214,049 propagules of 47 species), and a rough proxy of species abundance across the entire pool. We used both species and phylogenetically independent contrasts as data points, and accounted for variation in size of generative propagules. Results Both hypotheses were confirmed. Numbers of generative and vegetative propagules trapped per species were significantly negatively correlated (r = ?0.33; t₄₅ = ?2.61, P = 0.006) and thus strongly deviated from the null expectation. This was confirmed by analyses accounting for variation in species abundance across the species pool, and in the size of generative propagules. Main conclusions The results indicate that plant recruitment and community assembly across streams may be influenced by the way individual plants allocate their resources between competing life‐history functions. Life‐history evolution across angiosperms might thus have constrained the present‐day assembly of local communities.     

Interactions among species with contrasting dispersal modes explain distributions for epiphytic lichens

Understanding how the biodiversity response to climate change will be modified at ecological scales, e.g. by species interactions, is a major challenge. Lichen epiphytes – the close interdependent relationship between a heterotrophic fungus and photosynthetic partner (photobiont) – are used here to explore how interaction regimes (between lichen species, and between lichens and their photobionts) explain distribution patterns along spatial climatic gradients. To do this we tested field evidence for the ‘core‐fringe hypothesis’, which proposes a facilitative interaction; sexually‐reproducing and spore‐dispersed lichens with a requirement for resynthesis with a compatible photobiont (Nostoc) are facilitated by the prior establishment of asexual lichens which disperse both the fungus and photobiont together. We used two closely related Nephroma species which differ in their reproductive mode – N. laevigatum (sexual spore‐dispersed) and N. parile (asexual) – and compared their occurrence along a bioclimatic gradient to local habitat factors, including the co‐occurrence of asexual lichens which have shared specificity for compatible Nostoc genotypes. The results showed that: 1) N. laevigatum is significantly more likely to occur on trees that have already been colonised by asexual lichens with shared specificity for Nostoc, supporting the core‐fringe hypothesis, while 2) N. parile is independent of this association (strengthening the core‐fringe hypothesis), with its response to a precipitation gradient modified by microhabitat factors. This positive test for the core‐fringe hypothesis demonstrates how interaction regimes can fundamentally alter expectations under climate change. There is an assumption that spore‐dispersed lichen species could more easily track their suitable bioclimatic space through fragmented habitat, compared to asexual species with larger and heavier propagules. However, the establishment of spore‐dispersed lichen epiphytes into new habitat may be limited by the dispersal rates of asexual species, which act as key facilitators.     

Insulin-like growth factor-1 is associated with life-history variation across Mammalia

Despite the diversity of mammalian life histories, persistent patterns of covariation have been identified, such as the ‘fast–slow’ axis of life-history covariation. Smaller species generally exhibit ‘faster’ life histories, developing and reproducing rapidly, but dying young. Hormonal mechanisms with pleiotropic effects may mediate such broad patterns of life-history variation. Insulin-like growth factor 1 (IGF-1) is one such mechanism because heightened IGF-1 activity is related to traits associated with faster life histories, such as increased growth and reproduction, but decreased lifespan. Using comparative methods, we show that among 41 mammalian species, increased plasma IGF-1 concentrations are associated with fast life histories and altricial reproductive patterns. Interspecific path analyses show that the effects of IGF-1 on these broad patterns of life-history variation are through its direct effects on some individual life-history traits (adult body size, growth rate, basal metabolic rate) and through its indirect effects on the remaining life-history traits. Our results suggest that the role of IGF-1 as a mechanism mediating life-history variation is conserved over the evolutionary time period defining mammalian diversification, that hormone–trait linkages can evolve as a unit, and that suites of life-history traits could be adjusted in response to selection through changes in plasma IGF-1.     

Comparative demography of commercially important parrotfish species from Micronesia

Fishery‐independent sampling was used to determine growth patterns, life span, mortality rates and timing of maturation and sex change in 12 common parrotfishes (Labridae: tribe Scarinae) from five genera (Calotomus, Cetoscarus, Chlorurus, Hipposcarus and Scarus) in Micronesia. Interspecific variation in life‐history traits was explored using multivariate analysis. All species displayed strong sex‐specific patterns of length‐at‐age among which males reached larger asymptotic lengths. There was a high level of correlation among life‐history traits across species. Relationships between length‐based and age‐based variables were weakest, with a tenuous link between maximum body size and life span. Cluster analysis based on similarities among life‐history traits demonstrated that species were significantly grouped at two major levels. The first grouping was driven by length‐based variables (lengths at maturity and sex change and maximum length) and separated the small‐ and large‐bodied species. Within these, species were grouped by age‐based variables (age at maturity, mortality and life span). Groupings based on demographic and life‐history features were independent of phylogenetic relationships at the given taxonomic level. The results reiterate that body size is an important characteristic differentiating species, but interspecific variation in age‐based traits complicates its use as a life‐history proxy. Detailed life‐history metrics should facilitate future quantitative assessments of vulnerability to overexploitation in multispecies fisheries.     

Enzyme polymorphism inPreissia quadrata (Hepaticae, Marchantiaceae)

Preissia quadrata has a high habitat specificity and a reproductive system including frequent sexual reproduction and absence of specialized asexual propagules. Fertilization is aquatic. Colonies collected in Europe, Asia, eastern and western Canada show the species to be polymorphic at the eleven isozyme loci studied. Most often, thalli of the same colony appeared genetically identical. Partitioning of genetic variation was among rather than within colonies, suggesting a clonal structure of the colonies and a species structure consisting of a series of small populations reproductively isolated from each other. Little genetic exchange between colonies growing nearby was present, but the species does not appear to consist of regionally circumscribed genetically cohesive entities. Some European and Canadian colonies had identical electrophoretic patterns. The problem of understanding the phenomena holding liverwort species together, both morphologically and genetically, remains an open question. The genetic structure of the species might be due to its past history. It might have had a continuous distribution in the northern part of the Northern Hemisphere. If so, little subsequent genetic diversification, perhaps linked to its haploid-dominant life cycle, must be supposed.     

Developmental analysis of the evolutionary origin of vegetative propagules in Mimulus gemmiparus (Scrophulariaceae)

Mimulus gemmiparus (Scrophulariaceae), a rare endemic of Colorado, has a novel life history that depends on an unusual method of vegetative reproduction. The plants are functionally annuals; however, reproduction is asexual via propagules that have been termed gemmae. The morphological identity and the evolutionary antecedent of these propagules are unclear. We approached this problem through comparative developmental analyses of M. gemmiparus and the presumed progenitor species, Mimulus guttatus. In M. gemmiparus there are two meristems initiated in the axil of each leaf primordium. The distal meristem has the potential to produce either a lateral branch or a flower, and the proximal meristem becomes a vegetative propagule (the gemma) that is ultimately surrounded by an expanded, ensheathing petiole. The first leaves of the propagules are thickened and are the site of nutrient storage. Consequently, these propagules can be characterized morphologically as brood bulbils. Mimulus guttatus also has two meristems in each leaf axil; however, the proximal meristem typically remains dormant and serves no function in the life history of this species. Based on architectural and developmental correspondence, we hypothesize that the propagule of M. gemmiparus is homologous to the proximal meristem of M. guttatus. Comparative analysis shows that evolution of the bulbil has involved both the incorporation of features present in shoots of M. guttatus and the acquisition of novel features.     

A revision of Calvoa Hook. f. (Melastomataceae)

A revision of the genus Calvoa Hook. f. (Melastomataceae) is presented. Principal Component Analyses (PCA) and Cluster Analyses (UPGMA) were performed to elucidate three cases of difficult species delimitation and infraspecific variation, with the following results. (1) The separation between C. crassinoda Hook. f. and C. grandifolia Cogn. was clear in the results obtained with both methods. (2) The group of the West-Central African species Calvoa angolensis A. Fern. & R. Fern., C. calliantha Jacq.-Fél., C. monticola A. Chev., C. pulcherrima Gilg ex Engl., C. sapinii De Wild., C. seretii De Wild., C. sinuata Hook. f. and C. subquinquenervia De Wild, proved difficult to resolve with either analysis, but the Cluster Analysis produced results which are more consistent with the identification of the specimens. (3) Neither of the two analyses supported the recognition of infraspecific categories in Calvoa orientalis Taub. Eighteen species are recognized in the genus. The new species Calvoa Jacques-felixii E. Figueiredo is described and the new combination Calvoa seretii subsp. wildemaniana (Exell) E. Figueiredo is made. Four lectotypes are designated. The conservation status of some taxa is discussed. Six species are considered rare and possibly under threat.     

At the north-eastern extremity: variation in <Emphasis Type="Italic">Cepaea nemoralis</Emphasis> around Gdańsk,northern Poland

Variation in the shell colour and banding polymorphism in the land snail Cepaea nemoralis was studied in 260 populations in the region of Gdańsk, northern Poland. Unlike in other regions of Poland, many populations contain brown shells. Populations from shaded habitats have higher frequencies of brown than those from open and intermediate habitats, largely at the expense of yellow shells. Nearly all brown shells are also unbanded. Apart from this disequilibrium, banding morphs among yellow and pink shells show no relationship to habitat. There are no broad geographical trends in morph-frequencies, but there are very strong correlations among populations very close together, revealed both by pairwise analysis and Moran’s I. Principal Component Analyses show that these correlations relate to overall genetic similarity at the loci involved. The populations are at the north-eastern limits of the species’ range; habitats are mostly anthropogenic, and comparisons with studies in two urban areas (Wrocław, SW Poland, and Sheffield, central England) suggest that the patterns of variation seen are a product of human transport of propagules followed by local dispersal. The effect of habitat here is much less marked than in regions much further west, but it indicates that natural selection has occurred.     

CLASSIFICATION OF DIVERSE CALL TYPES USING CLUSTER ANALYSIS TECHNIQUES

ABSTRACT

We investigated the problem of categorizing the repertoire of a group of highly varied vocalizations. A set of Weddell seal Leptonychotes weddellim-air calls recorded near Davis, Antarctica, were examined. The repertoire size was estimated by first subjectively assigning each call (based on auditory and spectrographic patterns) to one of a large number of provisional call types. A set of frequency, duration and waveform measurements were made on every call. For each provisional call type, the mean value of each of these measures was calculated. These mean values were used to perform Cluster Analyses of the provisional call types. Beginning with calls clustered closest together, the most similar provisional call types were successively amalgamated until further joining would link two clearly dissimilar vocalizations (rising versus falling frequency sweeps in this case). The Weddell seal repertoire contained 12 call types ranging from long sinusoidal upsweeps to growls. This procedure provides a method of estimating the minimum repertoire size of a sample of calls. Dichotomous Sorting using Principal Components Analysis can also be used to categorize calls but will probably be of greater value when applied to finding subdivisions within a single call type.     

The macro-invertebrate communities of temporary pools in an intermittent stream in tropical Queensland

Nine pools were studied in an intermittent stream. They were close to each other but differed in size, substratum, amount of shade and duration of water presence. Initial conditions were set by wet season flooding, following which each pool developed its own character. Cluster analysis showed that communities changed substantially with time in both species composition and abundance, and that some initially similar communities diverged while some initially dissimilar ones converged.Principal Components Analysis showed that these community changes were due largely to the drying of the pools, and to seasonal effects. Specialised drying communities developed in the pools of longer duration (up to 8 months), but not in the short-lived pools (3–4 months). Apparently similar numerical responses in different pools were often caused by different suites of predominant species.The diversity of the fauna (total of 78 species) was high in comparison with similar habitats in non-tropical regions.The pattern of development of the pools' communities was much as expected from island biogeography theory, except that there was no relationship between species richness and duration and size of pools. The species complements were largely opportunistic, being governed by extreme environmental disturbance.     

Asexual propagules as an adaptive trait for epiphylly in tropical leafy liverworts (Lejeuneaceae)

? Premise of the study: Many links between form and function are described in the context of adaptation. Several morphological and life-history traits in the leafy liverwort family Lejeuneaceae (Marchantiophyta) have been hypothesized to be adaptations for living on the surface of leaves of vascular plants (epiphylly). There have been, however, no rigorous tests of these hypotheses. ? Methods: Using a recently published phylogeny of Lejeuneaceae and trait data from published monographs, I tested the correlations of putative adaptive traits with the incidence of epiphylly. Both cross-species and phylogenetic-based analyses of trait data were performed to distinguish the patterns of shared evolutionary history from independent origins of putatively adaptive traits. The rates of transitions between different combinations of character states were also calculated to determine whether traits were more likely to evolve in the presence of epiphylly. ? Key results: Only one trait, production of asexual propagules, was correlated with epiphylly in the phylogenetic-based analysis. The rate of transition to asexual propagules was also significantly higher in the presence of epiphylly. Other traits correlated with epiphylly appeared to be the results of shared evolutionary history among sister taxa and therefore not due to adaptive evolution. ? Conclusion: The present study distinguished production of asexual propagules from other traits as a key adaptive response to living on the leaf surface. No other putative "adaptive" traits to epiphylly showed evidence of being specific adaptation to epiphylly. The results highlight the importance of phylogenetically controlled methods in determining an adaptive function of traits.     

Evolution of Rotifer Life Histories

When compared to most other multicellular animals, rotifers are all relatively small, short-lived and fast-reproducing organisms. However among and within different rotifer species there is a large variation in life history patterns. This review accounts for such variation in rotifers, with a strong focus on monogonont rotifers. As the life cycle of monogonont rotifers involves both asexual and sexual reproduction, life history patterns can be examined on the level of the genetic individual, which includes all asexual females, sexual females and males that originated from one resting egg. This concept has been applied successfully in many areas, for example in predicting optimal levels of mictic reproduction or sex allocation theory. The benefits and implications of the view of the genetic individual are discussed in detail. Rotifer life histories can also be viewed on the level of physiological individuals. A large part of this review deals with the life histories of individual amictic females and addresses life history traits like body size, egg size and resource allocation patterns. It asks which trade-offs exist among those traits, how these traits change under the influence of environmental factors like food availability or temperature, and whether these changes can be interpreted as adaptive.     

A method for subgrouping the S-type of CMS forms in maize

Summary Simple correlations were calculated between nine different cms sources (cms-S, -R, -ML, -L, -CA, -EK, -C, -Rb, -T) on the basis of the weighted restoring reactions of 41 inbred lines. The Principal Component Analysis was applied to a 9 × 9 matrix which clearly grouped cytoplasms according to their similarities. The Principal Component I included S, R, ML, L, CA and EK cytoplasms; the Principal Component II contained C and Rb cytoplasms while T-cytoplasms was placed in Principal Component III. This corresponds to the main groupings indicated in the literature (Beckett 1971). However, after varimax rotation of the Principal Components, the S main group, including the 6 tested cytoplasms, fell into 3 subgroups: I.: S, R, ML; II.: L, CA; III.: EK.These data indicate that the Principal Component Analysis can be used to select a limited number of cms sources from the S group, representing the variability of the cytoplasmic gene pool of that group.     

Life history tactics of darters (Percidae: Etheostomatiini) and their relationship with body size, reproductive behaviour, latitude and rarity

Patterns of covariation of life history traits of darters in the genus Etheostoma are reviewed. The primary pattern is associated with body size. Large darters grow faster, mature at a larger size, produce bigger clutches, and have longer reproductive and life spans, and shorter spawning seasons, than do small darters. When the effects of size are removed statistically, the dominant secondary pattern matched the r-K continuum from fast-growing, short-lived, primarily semelparous species with many small ova and a high reproductive effort (r-species) to slow-growing, long-lived, iteroparous species with few large ova and a low reproductive effort (K-species). Variation in life history traits is also influenced by reproductive behaviour, latitude, and rarity (as measured by geographic range). There are significant differences in the primary and secondary life history patterns among reproductive guilds. Latitude and rarity are not correlated with these primary and secondary patterns. Instead, they account for variation of tertiary patterns. Rare species may not match the reproductive performance of more common and widely distributed species. Future studies of life history traits in darters should focus on species whose reproductive behaviour differs from that of the species reviewed in this study, and on the demographic characteristics of rare or declining populations.     

Soil fungi invest into asexual sporulation under resource scarcity,but trait spaces of individual isolates are unique

During the last few decades, a plethora of sequencing studies provided insight into fungal community composition under various environmental conditions. Still, the mechanisms of species assembly and fungal spread in soil remain largely unknown. While mycelial growth patterns are studied extensively, the abundant formation of asexual spores is often overlooked, though representing a substantial part of the fungal life cycle relevant for survival and dispersal. Here, we explore asexual sporulation (spore abundance, size and shape) in 32 co-occurring soil fungal isolates under varying resource conditions, to answer the question whether resource limitation triggers or inhibits fungal investment into reproduction. We further hypothesized that trade-offs exist in fungal investment towards growth, spore production and size. The results revealed overall increased fungal investment into spore production under resource limitations; however, effect sizes and response types varied strongly among fungal isolates. Such isolate-specific effects were apparent in all measured traits, resulting in unique trait spaces of individual isolates. This comprehensive dataset also elucidated variability in sporulation strategies and trade-offs with fungal growth and reproduction under resource scarcity, as only predicted by theoretical models before. The observed isolate-specific strategies likely underpin mechanisms of co-existence in this diverse group of saprobic soil fungi.     

Studies on Timmiella barbuloides (Brid.) Moenk., IV. SEM and TEM characterization of spore wall and first germination stages

Abstract

The spore wall morphology of Timmiella barbuloides (Pottiales, Musci) is described. The spores are catalept, with an ornamentation pattern consisting of unevenly spaced, shortly pedunculated pilum-and gemma-like processes. The spore coat consists of three, unevenly thick layers: intine, exine, and perine. The exine is not involved in wall ornamentation, the processes consisting of perine only. The leptoma, a spore coat area involved in germination, consists of an intine markedly thickening in an area of thinning exine and, outside, with a spore coat area where perinous processes become sparse. On the basis of observations and of the data reported in recent literature the classical definition of the leptoma is modified. It is considered to be a structurally specialized, but not necessarily thin, area.     

Dispersal potential of spores and asexual propagules in the epixylic hepatic Anastrophyllum hellerianum

Dispersal ability is of great importance for plants, which commonly occupy spatially and temporally limited substrate patches. Mixed reproductive strategies with abundant diaspore production are favoured in a heterogeneous landscape to ensure successful colonisation at different distances. In bryophytes, long-distance dispersal has been thought to take place primarily by spores, while asexual propagules are important in local dispersal and in the maintenance of colonies. In the present study, we investigated the dispersal potential of two equally sized propagules, sexually formed spores and asexually produced gemmae in the dioecious, epixylic hepatic, Anastrophyllum hellerianum, which inhabits spatially and temporally limited substrate patches. We trapped propagules at different distances (0–10 m) and directions from the source colonies in two experiments: one in a natural habitat within a forest and another involving an artificial set-up in an open habitat. Spore dispersal showed only slight distance dependence both in the open and the forest habitats, presumably as a consequence of wind affecting the dispersal pattern. Gemma dispersal was more strongly distance-dependent in the open habitat than in the forest sites. Considerably more gemmae were deposited during rainy than dry periods, possibly because of the effect of rain drops on gemma release. However, weather conditions had no effect on the dispersal patterns of spores or gemmae. In A. hellerianum, the combination of occasional spore production and practically continuous, massive gemma production facilitates dispersal both on local scale and over long distances. Unlike previously assumed, not only spores but also the asexual propagules may contribute to long-distance dispersal, thus allowing considerable gene flow at the landscape level.     

Geographic variation in the life history of the sagebrush lizard: the role of thermal constraints on activity

Thermal constraints on the time available for activity have been proposed as a proximate mechanism to explain variation in suites of life history traits. The longer that an ectotherm can maintain activity, the more time it has to forage and the greater chance that it will encounter a predator and be eaten. Thus, the thermal environment may produce a trade off between growth and survival when variation in the environment favors increased activity. I used mark-recapture data from a demographic study of three natural populations of the sagebrush lizard (Sceloporus graciosus) and estimates of thermal opportunity for each population to evaluate whether variation in the thermal environment can explain patterns of growth and survival that occur over an elevational gradient. Lizards from the highest elevation population exhibited higher individual growth rates than those of lizards from lower elevation, while mortality rates increased with elevation for these populations. The covariation of fast growth and high mortality with increased thermal opportunity is the opposite trend expected if the thermal environment alone is to explain patterns of life history in these lizards. Additional factors including thermal heterogeneity in the distribution of microhabitats of lizards, adaptation to local environmental conditions, and a potential trade-off between resource acquisition and predation risk need to be addressed to obtain a satisfactory explanation of the causative mechanisms producing life history variation.