Resource allocation patterns in a forb and a sedge in two arctic environments—short-term response to herbivory

The present work investigates C and N allocation patterns in two forage plants: a forb, Oxyria digyna , and a sedge, Eriophorum angustifolium , in subarctic Sweden and high arctic Canada. Short-term changes in concentrations after simulated or natural herbivory (caused by Gynaephora groenlandica on Oxyria in the high arctic habitat) were also investigated. There were no clear differences in concentrations of C and N between the high arctic and subarctic sites in either species. In Oxyria of the subarctic habitat, the minimum N concentrations occurred at earlier phenological stages compared with plants in the high arctic habitat. Simulated herbivory increased the concentration of C in belowground tissues relative to those in control plants in Oxyria at the subarctic site, which may indicate increased allocation of non-C compounds to the growing shoots or daughter ramets. Herbivory by Gynaephora groenlandica caterpillars increased the N concentrations of Oxyria both in aboveground and belowground tissues, possible indicating increased uptake of N in the high arctic habitat. Eriophorum did not show clear trends in concentrations relating to habitat, phenology or simulated herbivory. The difference between Oxyria and Eriophorum in their response to herbivory apparently resulted from contrasting growth habits between the species. Tiller death after reproduction and long leaf life span may be the main reasons for the lack of clear patterns in concentrations in Eriophorum. Compensation after herbivory may be attained by the early production of daughter tillers in Eriophorum instead of the regrowth of the damaged ramets, as in Oxyria. Monitoring the responses for only one season is apparently a too short time period in these long-lived plants.     

Nanoplanktonic coccolithophorids (Prymnesiophyceae, Haptophyceae) from the Weddell Sea, Antarctica

The examination of whole mounts prepared for transmission electron microscopy has resulted in the finding of thirteen taxa of nanoplanktonic coccolithophorids from the Weddell Sea, Antarctica. The material was collected as part of the AMERIEZ programme, March 1986. Cold-water adapted nanoplanktonic coccolithophorids have previously been shown to constitute a recurrent plankton element at subarctic and arctic localities. Three of the Weddell Sea species, Wigwamma annulifera, W. arctica , and Papposphaera sagittifera , are conspecific with northern hemisphere material, while two species, Calciarcus alaskensis and Turrisphaera arctica , are possibly identical with previously described arctidsubarctic material. Six taxa new to science have been described from the Weddell Sea, Wigwamma antarctica, W. triradiata, Trigonaspis melvillea, Pappomonas weddellensis, Papposphaera obpyramidalis , and P. simplicissima . The cooccurrence of identical forms at the two poles, and the fact that the species described are allocated to "arctic" genera, indicate a geologically relatively recent exchange of biological material between the poles.     

Effects of experimental warming on arctic willows (Salix spp.): a comparison of responses from the Canadian High Arctic, Alaskan Arctic, and Swedish Subarctic

Three species of dwarf, prostrate willow ( Salix arctica , S. rotundifolia and S. herbacea ) were subjected to experimental summer warming in high arctic Canada, arctic Alaska, and subarctic Sweden, respectively, as part of the International Tundra Experiment. Phenological and growth responses of these species were compared for the second season of the experiment. Stigmas became receptive and pollen dispersal occurred significantly earlier for S. rotundifolia and S. herbacea in the ITEX open-top chambers, but not for S. arctica . Warming had no effect on the timing of seed dispersal, leaf yellowing, or leaf senescence. The length and dry weight of the largest leaves were greater for warmed plants, and was significant for S. rotundifolia . The number of catkins/plot did not differ among species or treatments, but the fruit : flower ratio was reduced in the experimental plots.     

Further observations on Potamodiscus Gerloff =Pinaciophora Greeff, with special reference to Alaska and arctic Canada

Pinaciophora spiculata sp. nov. is described on the basis of two complete periplasts collected in arctic Canada at—l°C in 1973. One has been tested for silicon by means of X-ray microanalysis with strongly positive results. The species possesses dimorphic scales, the spicular scales and exceptionally small plate-scales being equally diagnostic. The circumstances under which the name has already appeared in the literature are explained. In addition, the known ranges of two other taxa are extended by broken periplasts recovered from other arctic or subarctic areas, notably P. denticulata Thomsen (plate scales only) formerly known from Peru and now from South Alaska (at Homer) while the type species of the genus ( Poturnodiscus kalbei Gerloff 1968= Pinaciophora fluivatilis Greeff 1875) is demonstrated both from South Alaska and from beneath sea ice in North Alaska, in both localities possibly introduced into the sea by freshwater drainage.     

Variability of macroinvertebrate community composition in an arctic and subarctic stream

The macroinvertebrate community composition was compared in two Alaskan streams (USA) for numeric and species constancy during the ice-free period from 1981 to 1983. Imnavait Creek is a first order arctic stream (60° 39 N, 149° 21 W) draining upland tundra in the foothills of the Brooks Range. Caribou-Poker Creek is a 4th order subarctic stream (65° 08 N, 147° 28 W) draining the taiga forest north of Fairbanks, Alaska. The aquatic insect larvae and other macroinvertebrates were sampled with drift nets and Hess bottom samplers for four periods, each 1 week long in the ice free season of three years. We found 112 species in the arctic stream and 138 species in the subarctic stream in a chironomid-dominated community. In any sample period the communities contained 51–60 species in the arctic and 49–92 species in the subarctic. Between the four sample periods on average 39% and 50% of the species were present in two sequential samples in the arctic and subarctic stream, respectively. New immigrants, never before found in the system, averaged 37% and 31% of the community, respectively. These systems are exposed to several intermediate disturbances: prolonged and variable freeze-up, extreme variation in discharge, wide diel and seasonal changes in temperature, and erosion by frazil and anchor ice. The dipterans that compose the most numerous and variable taxa must have variable diapause, ability to grow in cold waters, and good dispersal powers, even migrating across drainages in the arctic. Much of the seasonal dominance pattern appears therefore to be stochastic.     

Congeneric amphipods show differing abilities to maintain metabolic rates with latitude

Metabolic variability across latitudinal populations of gammarid amphipods was examined in the summer by determining whole-animal rates of oxygen uptake (M(o)?) in four species with overlapping distribution patterns in the northeast Atlantic and Arctic oceans. Comparisons were made between an arctic/boreal species, Gammarus setosus, a subarctic/boreal species, Gammarus oceanicus, a boreal/temperate species, Gammarus duebeni duebeni, and a temperate species, Gammarus locusta. Measurements included acclimatized M(o)? in all four species and M(o)? after acclimation to 10°C in two populations of G. oceanicus and G. locusta. In G. oceanicus, acclimatized M(o)? declined with latitude (13° to 5°C) so that metabolic rates were lower in subarctic (79°N) relative to temperate (58°N) populations and similar to the values in G. setosus at 79°N. Consequently, there was no evidence for metabolic rate compensation in the colder-water, high-latitude populations in the summer. Further examination of the specific effects of temperature revealed similarities in M(o)? between populations of G. oceanicus acclimated at 10°C and similarities in thermal sensitivity (Q(10)) and activation energies (E(a)) on exposure to acute temperature change. In sharp contrast, there was no variation in summer acclimatized M(o)? with latitude in either G. d. duebeni between 48° and 70°N or G. locusta between 38° and 53°N. Instead, the two species maintained relatively high metabolic rates across latitudes, which were associated in G. locusta with differences in M(o)? and with Q(10) and E(a) values in amphipods acclimated at 10°C. The ability to compensate metabolic rate with latitude in the summer suggests greater metabolic flexibility, which predicts a greater capacity for survival during climate change of the temperate/boreal over the subarctic and arctic gammarid species.     

AN SEM‐BASED ANALYSIS OF THE MORPHOLOGY,ANATOMY, AND REPRODUCTION OF LITHOTHAMNION TOPHIFORME (ESPER) UNGER (CORALLINALES,RHODOPHYTA), WITH A COMPARATIVE STUDY OF ASSOCIATED NORTH ATLANTIC ARCTIC/SUBARCTIC MELOBESIOIDEAE1

Lithothamnion tophiforme (Esper) Unger is a dominant, arctic, saxicolous species that extends southward, albeit with reduced cover, into the deeper colder waters of the North Atlantic subarctic, where it also occurs in significant rhodolith deposits with L. glaciale. The external appearance of L. tophiforme is distinctive, but typification, anatomy, reproduction, ecology, and biogeography have not been previously analyzed. These topics are now addressed, with extensive use of SEM, in comparison with other North Atlantic arctic and subarctic melobesioid genera and species. The species considered in this article comprise 95% of the coralline biomass of the colder North Atlantic and adjacent arctic (i.e. less than 12° C summer and less than 0° C winter). In the outer thallus region of coralline algae, crust extension proceeds, calcification develops, surface sloughing and grazing occur, and reproductive structures are initiated. Analysis of the ultrastructure of the outer thallus region (epithallium, meristem, and perithallium) of L. tophiforme shows distinctive generic similarities and specific differences from the other Lithothamnion species discussed here. Considerable generic differences from the Clathromorpum and Leptophytum species also encountered in the region considered are highlighted as well. We discuss the functional and taxonomic implications of these distinguishing features and recommend that they be more widely considered in future research on coralline algae to understand more fully the ecology and evolution of the Corallinales.     

Cortinarius subgenus Dermocybe in Greenland

The following species of Cortinarius subg. Dermocybe are recorded from Greenland: C. cinnamomeus, C. croceus, C. norvegicus, C. croceoconus, C. huronensis var. huronensis, C. uliginosus, C. cinnamomeoluteus , and C. polaris . Ecological and plant geographical features are discussed for each species. Cortinarius cinnamomeus and C. uliginosirs seem to be southern species never penetrating into the arctic zones; C. croceus, C. croceoconus, C. huronensis var. huronensis , and C. cinnamomeoluteus southern species penetrating into the arctic zones; and C. norvegicus and C. polaris true arctic species. Cortinarius norvegicus is probably continental, while the other species are distributed in continental as well as oceanic areas.     

Genetic and environmental effects on morphology in clonal sedges in the Eurasian Arctic

We studied the variation in morphological characters of importance for resource acquisition and storage in 21 populations of four clonal sedge taxa in arctic Eurasia, Carex bigelowii, C. ensifolia subsp. arctisibirica, C. lugens, and C. stans, and the response to transplantation to a common garden in Troms?, Norway. The morphology of C. stans was distinct from the other three taxa, all of which belong to the C. bigelowii species complex. However, differences among populations within taxa were even greater than differences among taxa, and environmental variables explained 40-50% of the among-population variation in the morphological characters. Stomatal size decreased with temperature while stomatal density increased. Shoot height and leaf width were smaller at peak lemming population phase, while rhizome length was shorter at higher longitudes. Transplantation to a common garden affected stomatal density in all taxa, stomatal size and shoot height only in some taxa, while leaf width was not affected. We found a weak, but highly significant correlation between geographical, morphological, and genetic distances. We concluded that although genotypic differentiation in arctic rhizomatous Carex species is reflected in their morphology, they are also capable of plastic morphological responses to the environment and that these responses are specific for each taxon.     

Global change effects on the long‐term feeding ecology and contaminant exposures of East Greenland polar bears

Rapid climate changes are occurring in the Arctic, with substantial repercussions for arctic ecosystems. It is challenging to assess ecosystem changes in remote polar environments, but one successful approach has entailed monitoring the diets of upper trophic level consumers. Quantitative fatty acid signature analysis (QFASA) and fatty acid carbon isotope (δ¹³C‐FA) patterns were used to assess diets of East Greenland (EG) polar bears (Ursus maritimus) (n = 310) over the past three decades. QFASA‐generated diet estimates indicated that, on average, EG bears mainly consumed arctic ringed seals (47.5 ± 2.1%), migratory subarctic harp (30.6 ± 1.5%) and hooded (16.7 ± 1.3%) seals and rarely, if ever, consumed bearded seals, narwhals or walruses. Ringed seal consumption declined by 14%/decade over 28 years (90.1 ± 2.5% in 1984 to 33.9 ± 11.1% in 2011). Hooded seal consumption increased by 9.5%/decade (0.0 ± 0.0% in 1984 to 25.9 ± 9.1% in 2011). This increase may include harp seal, since hooded and harp seal FA signatures were not as well differentiated relative to other prey species. Declining δ¹³C‐FA ratios supported shifts from more nearshore/benthic/ice‐associated prey to more offshore/pelagic/open‐water‐associated prey, consistent with diet estimates. Increased hooded seal and decreased ringed seal consumption occurred during years when the North Atlantic Oscillation (NAO) was lower. Thus, periods with warmer temperatures and less sea ice were associated with more subarctic and less arctic seal species consumption. These changes in the relative abundance, accessibility, or distribution of arctic and subarctic marine mammals may have health consequences for EG polar bears. For example, the diet change resulted in consistently slower temporal declines in adipose levels of legacy persistent organic pollutants, as the subarctic seals have higher contaminant burdens than arctic seals. Overall, considerable changes are occurring in the EG marine ecosystem, with consequences for contaminant dynamics.     

Climate and vegetation in China IV. Distribution of tree species along the thermal gradient

The aim of this study was to examine correlations between vegetational zones and thermal climate on the basis of analysis of the thermal distribution of main-component trees in each zone. The whole and main thermal distribution ranges for 73 tree species were obtained from their distributions along the warmth index (WI) gradicnt. Cluster analysis using the WI values at the upper and lower limits of the main distribution ranges as parameters gave seven species groups. The geographical distributions of the species in each group were described. Moreover, the frequency distribution curves of the seven species groups were determined along the WI grandient, and four borderlines (lines I, II, III and IV) were found among them. The WI values at the lines I, II and IV were 50, 90, 175°C month, respectively, and corresponded to boundaries between the subarctic and cool-temperate zones (I), the cool-and warm-temperate zones (II) and the warmtemperated and subtropical/tropical zones (IV), respectively. Line III corresponded to a WI of 135°C month. The WI values at the forest limit and at the lower limit of the subarctic forest were estimated, and their averages were 15.2 and 49.6°C month, respectively. In conclusion, thermal climatic zones in China could be divided by the following WI values: alpine (arctic) zone, WI<15; subarctic zone, 15–50; cool-temperate zone, 50–90; warm-temperate zone, 90–175; subtropical/tropical zone >175°C month.     

The evolution and distribution of Calyptozetes sarekensis (Acari: Oribatei)

Calyptozetes sarekensis , an oribatid mite found predominantly, though not exclusively in lichens, inhabits regions within the arctic circle and subarctically between latitudes 60^o and 70^o (Swedish Lapland, Norway, Bear Island, Spitzbergen, Jan Mayen, Iceland, Greenland, northern Canada). In recent years, however, the species has been recorded from a number of montane sites in Great Britain and southern Europe. There would appear to be only two possible explanations for die discontinuous distribution of this mite. (1) The species has been transported from its main arctic and subarctic area of distribution to its southern montane outposts by some agent (driftwood, birds, wind), or (2) in Britain and southern Europe it represents a relict species of the Ice Age, left stranded on high ground after the final retreat of the ice. Evidence is presented in support of the second hypothesis. In addition, it is suggested that its presence on the different land masses of its main arctic and subarctic range, which are separated from each other by wide stretches of ocean, could be explained by die hypothesis that it is a very ancient species daring from the time Laurasia was in existence some 60 or more million years ago. The alternative hypothesis of transportation of the species between one or more of these areas, however, is by no means ruled out.     

The Freezing Response of an Arctic Cushion Plant, Saxifraga caespitosa L.: Acclimation, Freezing Tolerance and Ice Nucleation

Cold hardiness in actively growing plants of Saxifraga caespitosaL., an arctic and subarctic cushion plant, was examined. Plantscollected from subarctic and arctic sites were cultivated ina phytotron at temperatures of 3, 9, 12 and 21 °C undera 24-h photoperiod, and examined for freezing tolerance usingcontrolled freezing at a cooling rate of 3–4 °C eitherin air or in moist sand. Post-freezing injury was assessed byvisual inspection and with chlorophyll fluorescence, which appearedto be well suited for the evaluation of injury in Saxifragaleaves. Freezing of excised leaves in moist sand distinguishedwell among the various treatments, but the differences werepartly masked by significant supercooling when the tissue wasfrozen in air. Excised leaves, meristems, stem tissue and flowerssupercooled to –9 to –15 °C, but in rosettesand in intact plants ice nucleation was initiated at –4to –7 °C. The arctic plants tended to be more coldhardy than the subarctic plants, but in plants from both locationscold hardiness increased significantly with decreasing growthtemperature. Plants grown at 12 °C or less developed resistanceto freezing, and excised leaves of arctic Saxifraga grown at3 °C survived temperatures down to about –20 °C.Exposure to –3 °C temperature for up to 5 d did notsignificantly enhance the hardiness obtained at 3 °C. Whenwhole plants of arctic Saxifraga were frozen, with roots protectedfrom freezing, they survived –15 °C and –25°C when cultivated at 12 and 3 °C, respectively, althougha high percentage of the leaves were killed. The basal levelof freezing tolerance maintained in these plants throughoutperiods of active growth may have adaptive significance in subarcticand arctic environments. Saxifraga caespitosa L., arctic, chlorophyll fluorescence, cold acclimation, cushion plant, freezing stress, freezing tolerance, ice nucleation, supercooling     

Future Climate Change Will Favour Non-Specialist Mammals in the (Sub)Arctics

Arctic and subarctic (i.e., [sub]arctic) ecosystems are predicted to be particularly susceptible to climate change. The area of tundra is expected to decrease and temperate climates will extend further north, affecting species inhabiting northern environments. Consequently, species at high latitudes should be especially susceptible to climate change, likely experiencing significant range contractions. Contrary to these expectations, our modelling of species distributions suggests that predicted climate change up to 2080 will favour most mammals presently inhabiting (sub)arctic Europe. Assuming full dispersal ability, most species will benefit from climate change, except for a few cold-climate specialists. However, most resident species will contract their ranges if they are not able to track their climatic niches, but no species is predicted to go extinct. If climate would change far beyond current predictions, however, species might disappear. The reason for the relative stability of mammalian presence might be that arctic regions have experienced large climatic shifts in the past, filtering out sensitive and range-restricted taxa. We also provide evidence that for most (sub)arctic mammals it is not climate change per se that will threaten them, but possible constraints on their dispersal ability and changes in community composition. Such impacts of future changes in species communities should receive more attention in literature.     

Male territorial aggression and androgen modulation in high latitude populations of the Sooty, Passerella iliaca sinuosa, and Red Fox Sparrow, Passerella iliaca zaboria

The Fox Sparrows, Passerella iliaca, include multiple groups and subspecies distributed at several latitudes from the Alaskan arctic to the southwestern United States. As such, this species represents a potential model for investigating latitudinal variation in androgen secretion and aggressive territoriality in male passerines. Breeding male Fox Sparrows from two subspecies within two groups, the Sooty Fox Sparrow, P. i. sinuosa, and the Red Fox Sparrow, P. i. zaboria, were assessed for aggressive territoriality and androgen responsiveness at multiple latitudes in arctic and subarctic Alaska. Subarctic Sooty Fox Sparrows had higher circulating androgen levels in the early (8.54 ng/ml) versus mid–late breeding season (2.44 ng/ml). Males in the mid–late breeding season did not up-regulate androgen secretion in response to social challenge, but were aggressive and spent more time within 5 m of a decoy during a simulated territorial intrusion (STI) than early breeding males. Male subarctic Red Fox Sparrows had slightly higher circulating androgen levels (2.29 ng/ml) than arctic males (1.10 ng/ml) in the mid–late breeding season. However, androgen levels were not correlated with blood collection time after a social challenge in either group, suggesting that neither arctic nor subarctic males up-regulate androgen secretion during the mid–late breeding period. Arctic males spent more time within 5 m of a decoy and sang less than subarctic males during an STI in the mid–late breeding season. These findings demonstrate that the Fox Sparrow is a tractable model for investigating the latitudinal regulation of aggressive territoriality and androgen responsiveness in passerines.     

A genetic determination of cryptic species (Copepoda: Calanoida) and their postglacial biogeography in North America

Allozyme electrophoresis of Hesperodiaptomus arcticus (Marsh) sensu lato at subarctic Churchill, Canada revealed significant deviations from random mating in several populations, as shown by the complete absence of certain heterozygous types. A broad geographic survey of 20 enzyme loci in 1077 individuals from 27 ponds at 11 arctic and alpine sites west of Hudson Bay demonstrated that there were three genetically distinct types. Morphological analysis of genetically classified populations uncovered consistent morphological differences that had been ignored previously. These morphological characteristics were used to reclassify preserved museum and private specimens from additional sites from the entire North American distribution of H. arcticus sensu lato. A detailed study of two of the distinct types co-occurring at Churchill showed reproduction timing and habitat differences consistent with reproductive isolation. I concluded that there are three species and each have different distributions. Most regions examined must have been repopulated by dispersal from glacial refugia after deglaciation. The nominal species is apparently restricted to western arctic localities in the vicinity of the Alaskan refugium while the other two putative taxa have dispersed widely into previously glaciated regions.     

Phenotypic selection on floral traits in the arctic plant Parrya nudicaulis (Brassicaceae)

The evolution of floral traits is often attributed to pollinator‐mediated selection; however, the importance of pollinators as selective agents in arctic environments is poorly resolved. In arctic and subarctic regions that are thought to be pollen limited, selection is expected to either favor floral traits that increase pollinator attraction or promote reproductive assurance through selfing. We quantified phenotypic selection on floral traits in two arctic and two subarctic populations of the self‐compatible, but largely pollinator‐dependent, Parrya nudicaulis. Additionally, we measured selection in plants in both open pollination and pollen augmentation treatments to estimate selection imposed by pollinators in one population. Seed production was found to be limited by pollen availability and strong directional selection on flower number was observed. We did not detect consistently greater magnitudes of selection on floral traits in the arctic relative to the subarctic populations. Directional selection for more pigmented flowers in one arctic population was observed, however. In some populations, selection on flower color was found to interact with other traits. We did not detect consistently stronger selection gradients across all traits for plants exposed to pollinator selection relative to those in the pollen augmentation treatment; however, directional selection tended to be higher for some floral traits in open‐pollinated plants.     

Small changes in timing of breeding among subarctic passerines over a 32‐year period

Many bird populations in temperate regions have advanced their timing of breeding in response to a warming climate in recent decades. However, long‐term trends in temperature differ geographically and between seasons, and so do responses of local breeding populations. Data on breeding bird phenology from subarctic and arctic passerine populations are scarce, and relatively little data has been recorded in open‐nesting species. We investigated the timing of breeding and its relationship to spring temperature of 14 mainly open‐nesting passerine species in subarctic Swedish Lapland over a period of 32 years (1984–2015). We estimated timing of breeding from the progress of post‐juvenile moult in mist‐netted birds, a new method exploring the fact that the progress of post‐juvenile moult correlates with age. Although there was a numerical tendency for earlier breeding in most species (on average ?0.09 days/year), changes were statistically significant in only three species (by ?0.16 to ?0.23 days/year). These figures are relatively low compared with what has been found in other long‐term studies but are similar to a few other studies in subarctic areas. Generally, annual hatching dates were negatively correlated with mean temperature in May. This correlation was stronger in long‐distance than in short‐distance migrants. Although annual temperatures at high northern latitudes have increased over recent decades, there was no long‐term increase in mean temperature in May over the study period at this subarctic site. This is probably the main reason why there were only small long‐term changes in hatching dates.     

Calvatia sporocristata sp. nov. (Gasteromycetes) from Costa Rica

Calvatia sporocristata, Gasteromycetes, is described and illustrated as a new species. Its taxonomic characters are discussed and compared with other closely related species of the genus. Few studies have been carried out on tropical Gasteromycetes. The new species here proposed has, as the main features: exoperidium corky, smooth, one-layered, pseudoparenchymatous; endoperidium thin, papyraceous, dextrinoid; subgleba cellular; capillitium septate, dissociated at the septal site, lycoperdon type, without pores, elastic and spores amigdaloid, 4-6 (-7) x 2.5-4.5 microns with spines aligned, forming crests, 0.8 micron high.