Adaptation to marginal habitats by evolution of increased phenotypic plasticity

In an island population receiving immigrants from a larger continental population, gene flow causes maladaptation, decreasing mean fitness and producing continued directional selection to restore the local mean phenotype to its optimum. We show that this causes higher plasticity to evolve on the island than on the continent at migration-selection equilibrium, assuming genetic variation of reaction norms is such that phenotypic variance is higher on the island, where phenotypes are not canalized. For a species distributed continuously in space along an environmental gradient, higher plasticity evolves at the edges of the geographic range, and in environments where phenotypes are not canalized. Constant or evolving partially adaptive plasticity also alleviates maladaptation owing to gene flow in a heterogeneous environment and produces higher mean fitness and larger population size in marginal populations, preventing them from becoming sinks and facilitating invasion of new habitats. Our results shed light on the widely observed involvement of partially adaptive plasticity in phenotypic clines, and on the mechanisms causing geographic variation in plasticity.     

Species concept and morphological differentiation of strains traditionally assigned to Micrasterias truncata

The morphological and molecular differentiation of the Micrasterias truncata (Corda) ex Bréb species complex was investigated. In total, 17 strains traditionally assigned to M. truncata were isolated from different European localities (Czech Republic, southwest France, Ireland), and obtained from public culture collections. In addition, strains of the morphologically similar species, M. decemdentata (Nägeli) W. Archer and M. zeylanica F. E. Fritsch, were also included. Molecular phylogenetic analysis based on trnG^ucc intron sequences revealed five well supported clades. Two Australian strains assigned to M. truncata var. pusilla G. S. West formed a lineage sister to M. zeylanica. This was evident from a concatenated phylogeny based on small subunit rDNA and trnG^ucc intron sequences. The isolated position of these strains was also illustrated by parallel landmark‐based geometric morphometric analysis of cell shapes. The strains NIES 783 and NIES 784 probably represent a separate species. Particular analysis, including additional strains, is needed to resolve the relationship inside this lineage. The second phylogenetic lineage, containing two strains of M. truncata var. semiradiata (Kützing) Wolle, was also different from other strains on the basis of morphometric data. We suggest recognizing this variety as a separate species, Micrasterias semiradiata L.A. Brébisson ex F. T. Kützing. The remaining three clades formed a firmly supported group of the ‘core’M. truncata recognized by both molecular markers. However, neither any morphological, morphometric, nor geographical pattern was detected among members of these three clades. This pattern could be caused by a relatively recent origin of these lineages that may represent a sympatric, truly cryptic species. Strains attributable to traditional morphologically defined variety M. truncata var. neodamensis were nested within the ‘core’M. truncata.     

A climatic basis for microrefugia: the influence of terrain on climate

There is compelling evidence from glacial and interglacial periods of the Quaternary of the utilization of microrefugia. Microrefugia are sites that support locally favorable climates amidst unfavorable regional climates, which allow populations of species to persist outside of their main distributions. Knowledge of the location of microrefugia has important implications for climate change research as it will influence our understanding of the spatial distribution of species through time, their patterns of genetic diversity, and potential dispersal rates in response to climate shifts. Indeed, the implications of microrefugia are profound and yet we know surprisingly little about their climatic basis; what climatic processes can support their subsistence, where they may occur, their climatic traits, and the relevance of these locations for climate change research. Here I examine the climatic basis for microrefugia and assert that the interaction between regional advective influences and local terrain influences will define the distribution and nature of microrefugia. I review the climatic processes that can support their subsistence and from this climatic basis: (1) infer traits of the spatial distribution of microrefugia and how this may change through time; (2) review assertions about their landscape position and what it can tell us about regional climates; and (3) demonstrate an approach to forecasting where microrefugia may occur in the future. This synthesis highlights the importance of landscape physiography in shaping the adaptive response of biota to climate change.     

Impacts of climate warming on hybrid zone movement: Geographically diffuse and biologically porous “species borders”

Abstract The ecology and evolutionary biology of insect–plant associations has realized extensive attention, especially during the past 60 years. The classifications (categorical designations) of continuous variation in biodiversity, ranging from global patterns (e.g., latitudinal gradients in species richness/diversity and degree of herbivore feeding specialization) to localized insect–plant associations that span the biospectrum from polyphenisms, polymorphisms, biotypes, demes, host races, to cryptic species, remain academically contentious. Semantic and biosystematic (taxonomical) disagreements sometimes detract from more important ecological and evolutionary processes that drive diversification, the dynamics of gene flow and local extinctions. This review addresses several aspects of insect specialization, host‐associated divergence and ecological (including “hybrid”) speciation, with special reference to the climate warming impacts on species borders of hybridizing swallowtail butterflies (Papilionidae). Interspecific hybrid introgression may result in collapse of multi‐species communities or increase species numbers via homoploid hybrid speciation. We may see diverging, merging, or emerging genotypes across hybrid zones, all part of the ongoing processes of evolution. Molecular analyses of genetic mosaics and genomic dynamics with “divergence hitchhiking”, combined with ecological, ethological and physiological studies of “species porosity”, have already begun to unveil some answers for some important ecological/evolutionary questions. (i) How rapidly can host‐associated divergence lead to new species (and why doesn't it always do so, e.g., resulting in “incomplete” speciation)? (ii) How might “speciation genes” function, and how/where would we find them? (iii) Can oscillations from specialists to generalists and back to specialists help explain global diversity in herbivorous insects? (iv) How could recombinant interspecific hybridization lead to divergence and speciation? From ancient phytochemically defined angiosperm affiliations to recent and very local geographical mosaics, the Papilionidae (swallowtail butterflies) have provided a model for enhanced understanding of ecological patterns and evolutionary processes, including host‐associated genetic divergence, genomic mosaics, genetic hitchhiking and sex‐linked speciation genes. Apparent homoploid hybrid speciation in Papilio appears to have been catalyzed by climate warming‐induced interspecific introgression of some, but not all, species diagnostic traits, reflecting strong divergent selection (discordant), especially on the Z (= X) chromosome. Reproductive isolation of these novel recombinant hybrid genotypes appears to be accomplished via a delayed post‐diapause emergence or temporal isolation, and is perhaps aided by the thermal landscape. Changing thermal landscapes appear to have created (and may destroy) novel recombinant hybrid genotypes and hybrid species.     

Thermoregulation behaviour in codling moth larvae

Abstract. The thermoregulation behaviour of the codling moth, Cydia pomonella, is investigated in temperature gradient experiments with larvae feeding within apples, and with mature larvae searching for cocooning sites. Feeding larvae appear to prefer the apple hemisphere with a higher temperature (i.e. they build larger cavities in the radiated, warmer part of the fruit). The proportion of larval cavities in the warmer hemisphere is positively related to increasing apple temperature on that side, as well as to the temperature difference between the warm and the cool fruit hemisphere. The mechanism in feeding larvae can be termed as cryptic basking because, during microhabitat selection, the caterpillars exploit temperature differences that are caused explicitly by incident solar radiation. Fifth-instar larvae in search of cocooning sites show no temperature preference within the large gradient offered (9–29 °C), with no difference between males and females. During larval development, the insect changes its thermoregulation behaviour in response to a possible shift in benefits of an elevated body temperature with respect to environmental conditions. Both the thermoregulation behaviour and such a shift of behavioural response should be respected when simulating body temperatures of the species.     

Female genital system of the folding-trapdoor spider Antrodiaetus unicolor (Hentz, 1842) (Antrodiaetidae, Araneae): ultrastructural study of form and function with notes on reproductive biology of spiders

The genitalia of the female folding-trapdoor spider Antrodiaetus unicolor are characterized by two pairs of spermathecae that are arranged in a single row and connected to the roof of the bursa copulatrix. Each single spermatheca is divided into three main parts: stalk, bowl, and bulb, which are surrounded by the spermathecal gland. The epithelium of the spermathecal gland is underlain by a muscle meshwork and consists of different types of cells partly belonging to glandular cell units (Class 3 gland cells) that extend into pores in the cuticle of the stalk and bowl. Interestingly, the bulb lacks glandular pores and is characterized by a weakly sclerotized cuticle. This peculiarly structured bulb probably plays an important role in the discharge of the sperm mass. It is suggested that by contraction of the muscle layer the sperm mass may be squeezed out, when the bulb invaginates and expands into the spermathecal lumen, pushing the sperm to the uterus lumen. Each glandular unit consists of usually one or two central secretory cells that are for the most part surrounded by a connecting cell that again is surrounded by a canal cell. The canal cell, finally, is separated from the other epithelial cells (intercalary cells) located between the glandular units by several thin sheath cells that form the outer enveloping layer of the unit. The secretions are released through a cuticular duct that originates proximally between the apical part of the connecting cell and the apical microvilli of the secretory cells and runs into a pore of the spermathecal cuticle. The glandular products of the Class 3 gland cells likely contribute to the conditions allowing long-term storage of the spermatozoa in this species. Details regarding the ovary, the uterus internus, and the uterus externus are reported. Most of the secretion that composes the chorion of the egg is produced in the ovary. Glandular cell units observed in the uterus externus differ structurally from those in the spermathecae and likely play a different role. Finally, we briefly discuss our results on the female genitalia of A. unicolor in the light of knowledge about the reproductive biology of spiders.     

The benefits of polyandry in the free-spawning polychaete Galeolaria caespitosa

In many species, females are thought to benefit from polyandry due to the reduced risks of fertilization by genetically incompatible sperm. However, few studies that have reported such benefits have directly attributed variation in female reproductive success to the interacting effects of males and females at fertilization. In this paper, we determine whether male x female interactions influence fertilization in vitro in the free-spawning, sessile polychaete Galeolaria caespitosa. Furthermore, we determined whether polyandry results in direct fertilization benefits for females by experimentally manipulating the number of males contributing towards staged spawning events. To test for male x female interaction effects we performed an initial experiment that crossed seven males with six females (in all 42 combinations), enabling us to assess fertilization rates for each specific male-female pairing and attribute variation in fertilization success to males, females and their interaction. This initial experiment revealed a strong interaction between males and females at fertilization, confirming that certain male-female combinations were more compatible than others. A second experiment tested the hypothesis that polyandry enhances female reproductive success by exposing each female's eggs to either a single male's sperm (monandry) or the sperm from three males simultaneously (polyandry). We performed this second experiment at two ecologically relevant sperm concentrations. This latter experiment revealed a strong fertilization benefit of polyandry, independent of the effects of sperm concentration (which were also significant). We suggest that these direct fertilization gains arising from polyandry will constitute an important source of selection on females to mate multiply in nature.     

Phylogeography of a nematode (Heligmosomoides polygyrus) in the western Palearctic region: persistence of northern cryptic populations during ice ages?

This study establishes the continental phylogeographical pattern of a European nematode, Heligmosomoides polygyrus (Dujardin, 1845; Heligmosomoidea). We sequenced 687 base pairs of the mitochondrial DNA (mtDNA) cyt b gene for 136 individuals collected in 22 localities. The results revealed that H. polygyrus populations are separated into five major units corresponding to the Italian, northern European (Denmark and Ireland), Iberian, western European, and Balkan populations. Different subclades were also observed within the first two groups. Based on the rate of molecular evolution of H. polygyrus cyt b gene-estimated to 3.5%-3.7% divergence per million years (Myr) in a previous study--the isolation time of the five clades was estimated between 2.5 +/- 0.24 and 1.5 +/- 0.23 million years ago. Moreover, H. polygyrus presents a higher genetic variability in the Mediterranean peninsulas as compared to northwestern Europe, highlighting the role of these regions as refuge areas. Like its specific host, the wood mouse Apodemus sylvaticus, H. polygyrus' pattern of postglacial recolonization of northwestern Europe was initiated from Iberian populations, while Italian and Balkan populations did not expand to the north. The results also suggest the existence of forested and temperate refuges in the southern British Isles during the Quaternary. Finally, the genetic diversity as well as the level of genetic divergence between the lineages of H. polygyrus are compared to those observed in other vertebrate and invertebrate phylogeographical studies: the existence of highly differentiated lineages in H. polygyrus (5%-10% of genetic divergence) highlights that the effects of Pleistocene climate changes on free-living organisms are also reflected in their obligate parasites.     

Ribosomal DNA sequence variation among sympatric strains of the Cyclotella meneghiniana complex (Bacillariophyceae) reveals cryptic diversity

Cyclotella meneghiniana Kützing is one of the most commonly found and intensively studied freshwater diatom species. However, it is considered taxonomically problematic because of its unusually wide ecological range and large frustule ultrastructural variation. As part of a study of morphological and genetic variation in this morphospecies, we surveyed nucleotide variation in the hypervariable D1/D2 regions of the 28S rDNA, in the ribosomal internal transcribed spacer region (containing ITS1, the 5.8S rDNA and ITS2) and in the 18S rDNA in a collection of 20 sympatric strains. High genetic variability and strong indications of genetic structure among the Cyclotella meneghiniana strains were found. Representatives of four genetically distinct--apparently reproductively isolated--groups were revealed among them. The random distribution of ITS variation within these four groups indicated that the genetic structure in Cyclotella meneghiniana can probably be explained by the presence of cryptic sexual species rather than by the lack of allogamous sexual reproduction. The morphological features traditionally used for species identification in this group cannot distinguish these putative cryptic species.     

SPECIES ISOLATION,GENITAL MECHANICS,AND THE EVOLUTION OF SPECIES-SPECIFIC GENITALIA IN THREE SPECIES OF MACRODACTYLUS BEETLES (COLEOPTERA,SCARABEIDAE, MELOLONTHINAE)

The question asked was why male genitalic structures have diverged in three syntopic species of Macrodactylus beetles. Four hypotheses were evaluated: 1. The ways in which male genitalia mesh with internal female structures indicate that selection for species isolation via mechanical exclusion (“lock and key”) is unlikely to explain the genitalic differences. 2. The specific mate recognition hypothesis also clearly fails to explain genitalic differences due to the implausibility of postulated environmental effects on genitalia, and lack of postulated coevolution of male and female morphologies. 3. Selection for species isolation via differences in genitalic stimulation (sensory lock and key) is unlikely due to relatively infrequent cross-specific pair formation and intromission in the field, and “excessive” numbers of species-specific genitalic structures and male courtship behavior patterns which nevertheless occasionally fail. It also fails to explain the frequent failure of intraspecific copulations to result in sperm transfer. This hypothesis cannot, however, be rejected as confidently as the previous hypotheses. 4. Conditions under which sexual selection by cryptic female choice could take place are common. Females frequently exercise their ability to prevent sperm transfer by conspecific males even after intromission has occurred, and females generally mate repeatedly, probably with different males. Males behave as if cryptic female choice is occurring, courting assiduously while their genitalia are within the female. Sexual selection by female choice could thus contribute to the divergence in genitalic structures.