Phenology and life history of Belgica antarctica, an Antarctic midge (Diptera: Chironomidae)

ABSTRACT.

• 1 An account of the life-history with emphasis on phenology and number of instars is presented for Belgica antarctica Jacobs, the southernmost holometabolous insect.
• 2 Contrary to earlier reports, Belgica has four instars, in common with most other chironomids. Mean head capsule lengths varied between different populations but no overlap was found between discrete size classes of successive instars.
• 3 Belgica overwinters in all four instars.
• 4 Relative frequency of different instars from samples taken through the season indicates that a 2-year life-span is the commonest pattern.
• 5 Emergence of adults is relatively synchronous. Belgica exhibits protandry, which may be established at the time of pupation.

     

Aspects of the biology of the midge,Eretmoptera murphyi Schaeffer (Diptera: Chironomidae), introduced to Signy Island,maritime Antarctic

Summary The biology of a population of the subantarctic chironomid midge Eretmoptera murphyi Schaeffer, introduced to Signy Island in the maritime Antarctic more than 20 years ago, is described. Investment in reproduction by the parthenogenetic adult females is high, with individuals producing single egg batches containing ca. 85 eggs and having a dry weight of more than twice that of the spent female. In culture, egg development rates to hatching are increased significantly by increasing temperature from 2° to 12°C (a range covering mean summer temperatures found in the species' maritime Antarctic habitat, and natural habitat in the subantarctic). The gelatinous matrix of the egg batch forms a skin on drying, which may reduce further water loss, and allow the eggs or pre-emergence larvae to survive the short periods of desiccating conditions likely to occur in their natural habitat. The biology of E. murphyi is compared with that of the endemic maritime Antarctic species Belgica antarctica, showing much similarity. E. murphyi possesses several preadaptations which allow it to survive the harsher conditions of the maritime Antarctic.     

A molecular phylogeny of antarctic chironomidae and its implications for biogeographical history

The chironomid midges Belgica antarctica, Eretmoptera murphyi (subfamily Orthocladiinae) and Parochlus steinenii (subfamily Podonominae), are the only Diptera species currently found in Antarctica. The relationships between these species and a range of further taxa of Chironomidae were examined by sequencing domains 1 and 3–5 of 28S ribosomal RNA. The resulting molecular relationships between B. antarctica and E. murphyi, within Orthocladiinae, were highly supported by validation analyses, confirming their position within Chironomidae, as generated by classical taxonomy. Within Podonominae, P. steinenii from the Maritime Antarctic was more closely related to material from sub-Antarctic South Georgia than to material from Patagonia. Taking advantage of the availability of a molecular substitution rate calculated for this gene in Diptera, a dating of divergence between our study taxa was tentatively established. The divergence dates obtained were 49 million years (Myr), between B. antarctica and E. murphyi, and 68.5 Myr between these species and the closest Orthocladiinae taxon tested from Patagonia, suggesting that B. antarctica and E. murphyi were representatives of an ancient lineage. As both are endemic to their respective tectonic microplates, their contemporary distribution is, therefore, likely to have been shaped by vicariance rather than dispersal.     

Absence of metabolic cold adaptation and compensatory acclimation in the Antarctic fly, Belgica antarctica

The respiratory metabolism in larvae of the Antarctic fly, Belgica antarctica Jacobs (Diptera: Chironomidae) was investigated at Palmer Station, Anvers Island (64°46′S, 64°03′W). Oxygen consumption was linearly related to temperature from 0 to 20°C, respectively, 49 and 338 nl/mg live wt/hr. Maintenance at 0 and 10°C for 8 days had no differential effect on the metabolic rate, suggesting that larvae lack the ability for compensatory acclimation. A comparison of standard metabolism for polar and temperate chironomids revealed no elevation of metabolic rate in polar forms. However, polar species exhibited lower activation energies than temperate forms indicating that the respiratory metabolism of polar chironomids is relatively temperature independent.     

Antarctic hairgrass expansion in the South Shetland archipelago and Antarctic Peninsula revisited

Populations of both native higher Antarctic plants, Deschampsia antarctica and Colobanthus quitensis, increased during the last decades. However, for D. antarctica, previous population studies on the South Shetland Islands and the Antarctic Peninsula have been too sporadic, patchy, and methodologically different to allow general conclusions. Our aim was to compare sites with D. antarctica along a north–south latitudinal transect with an integral census method to assess the possible impact of climatic change on grass population dynamics. During two summer seasons (2009–2010), plant populations were censed on Fildes and Coppermine Peninsula and several localities on the west coast of the Antarctic Peninsula. Largest plant populations were found on Fildes Peninsula with vegetation cover (VC) of 44–46%. Six out of eleven stands of D. antarctica on Coppermine Peninsula were new records, with increasing plant number and VC (0.1–22%). In the Antarctic Peninsula, contrarily to our expectation, only at Forbes Point, D. antarctica VC was relatively high (ca. 2%) and a new stand of C. quitensis was found. At three previously reported sites, plants had disappeared. Our monitoring confirms that northern D. antarctica populations are expanding, but that this expansion is not continuous along the Antarctic Peninsula and inconsistent with the gradient of relative temperature increase in north–south direction. We suggest that other abiotic and biotic factors are influencing the colonization and expansion of vascular plants in this particular ecosystem.     

Function and immuno-localization of aquaporins in the Antarctic midge Belgica antarctica

Aquaporin (AQP) water channel proteins play key roles in water movement across cell membranes. Extending previous reports of cryoprotective functions in insects, this study examines roles of AQPs in response to dehydration, rehydration, and freezing, and their distribution in specific tissues of the Antarctic midge, Belgica antarctica (Diptera, Chironomidae). When AQPs were blocked using mercuric chloride, tissue dehydration tolerance increased in response to hypertonic challenge, and susceptibility to overhydration decreased in a hypotonic solution. Blocking AQPs decreased the ability of tissues from the midgut and Malpighian tubules to tolerate freezing, but only minimal changes were noted in cellular viability of the fat body. Immuno-localization revealed that a DRIP-like protein (a Drosophila aquaporin), AQP2- and AQP3 (aquaglyceroporin)-like proteins were present in most larval tissues. DRIP- and AQP2-like proteins were also present in the gut of adult midges, but AQP4-like protein was not detectable in any tissues we examined. Western blotting indicated that larval AQP2-like protein levels were increased in response to dehydration, rehydration and freezing, whereas, in adults DRIP-, AQP2-, and AQP3-like proteins were elevated by dehydration. These results imply a vital role for aquaporin/aquaglyceroporins in water relations and freezing tolerance in B. antarctica.     

A puzzle with many pieces: the genetic structure and diversity of Phaeocystis antarctica Karsten (Prymnesiophyta)

Strong ocean current systems characterize the Southern Ocean. The genetic structure of marine phytoplankton species is believed to depend mainly on currents. Genetic estimates of the relatedness of populations of phytoplankton species therefore should provide a proxy showing to what extent different geographic regions are interconnected by the ocean current systems. In this study, spatial and temporal patterns of genetic diversity were studied in the circumpolar prymnesiophyte Phaeocystis antarctica Karsten using seven nuclear microsatellite loci. Analyses were conducted for 86 P. antarctica isolates sampled around the Antarctic continent between 1982 and 2007. The results revealed high genetic diversity without single genotypes recurring even among isolates within a bloom or originating from the same bucket of water. Populations of P. antarctica were significantly differentiated among the oceanic regions. However, some geographically distant populations were more closely related to each other than they were to other geographically close populations. Temporal haplotype turnover within regions was also suggested by the multilocus fingerprints. Our data suggest that even within blooms of P. antarctica genetic diversity and population sizes are large but exchange between different regions can be limited. Positive and significant inbreeding coefficients hint at further regional substructure of populations, suggesting that patches, once isolated from one another, may not reconnect. These data emphasize that even for planktonic species in a marine ecosystem that is influenced by strong currents, significant breaks in gene flow may occur.     

Retrotransposon‐based genetic diversity of Deschampsia antarctica Desv. from King George Island (Maritime Antarctic)

Deschampsia antarctica Desv. can be found in diverse Antarctic habitats which may vary considerably in terms of environmental conditions and soil properties. As a result, the species is characterized by wide ecotypic variation in terms of both morphological and anatomical traits. The species is a unique example of an organism that can successfully colonize inhospitable regions due to its phenomenal ability to adapt to both the local mosaic of microhabitats and to general climatic fluctuations. For this reason, D. antarctica has been widely investigated in studies analyzing morphophysiological and biochemical responses to various abiotic stresses (frost, drought, salinity, increased UV radiation). However, there is little evidence to indicate whether the observed polymorphism is accompanied by the corresponding genetic variation. In the present study, retrotransposon‐based iPBS markers were used to trace the genetic variation of D. antarctica collected in nine sites of the Arctowski oasis on King George Island (Western Antarctic). The genotyping of 165 individuals from nine populations with seven iPBS primers revealed 125 amplification products, 15 of which (12%) were polymorphic, with an average of 5.6% polymorphic fragments per population. Only one of the polymorphic fragments, observed in population 6, was represented as a private band. The analyzed specimens were characterized by low genetic diversity (uH_e = 0.021, I = 0.030) and high population differentiation (F_ST = 0.4874). An analysis of Fu's F_S statistics and mismatch distribution in most populations (excluding population 2, 6 and 9) revealed demographic/spatial expansion, whereas significant traces of reduction in effective population size were found in three populations (1, 3 and 5). The iPBS markers revealed genetic polymorphism of D. antarctica, which could be attributed to the mobilization of random transposable elements, unique features of reproductive biology, and/or geographic location of the examined populations.     

Desmarestia antarctica (Desmarestiales,Phaeophyceae), a new ligulate Antarctic species with an endophytic gametophyte

Desmarestia antarctica Moe & Silva, a new species from the Antarctic Peninsula and the Scotia Arc, is distinguished from all other ligulate members of the genus by the arrangement of zoosporangia, which are produced together with paraphyses in a raised sorus. Among all species in the genus, onlyD. anceps Montagne shares this feature.Desmarestia antarctica is the first species of the genus for which an endophytic gametophyte is demonstrated, the usual host beingCurdiea racovitzae De Wildeman, a red alga of the familyGracilariaceae. The sporophyte contains only a moderate concentration of acid compared to Northern Hemisphere ligulate species, while physodes that probably contain polyphenolic compounds form noticeable speckles and dark margins in preserved plants. Often more than one axis arises from a single holdfast, probably as the result of compounding rather than proliferation.Dedicated to Prof. DrLothar Geitler on the occasion of the 90th anniversary of his birthday.     

Eretmoptera murphyi: pre‐adapted to survive a colder climate

During the late 1960s, larvae of the flightless midge Eretmoptera murphyi Schaeffer were accidentally transferred from the sub‐Antarctic island of South Georgia to Signy Island in the maritime Antarctic. Higher insects are rare in the Antarctic and the introduction and establishment of a new species is an unusual event. The fly has overcome the two major barriers to colonization of the Antarctic by new species: the geographical isolation of the region and its severe climate. Larvae of the flightless midge overwinter in the surface layers of soil on Signy Island where the temperature may fall to below ?10 °C, compared with as little as ?1.5 °C on South Georgia. This suggests the possession of a level of pre‐adaption to colder conditions. Summer‐collected larvae have a supercooling point (SCP or whole body freezing point) of approximately ?5.0 °C but survive experimental exposure to ?13 °C, giving them a level of freeze tolerance. After acclimation at ?4 °C for 4 days, the SCP changes little but the temperature at which 50% of the population would die decreases to lower than ?19 °C. Larvae are also resistant to dehydration. Under experimental conditions of 88% relative humidity at 5 °C, larvae lose water linearly (0.42% h^?1) over the first 30 h but resist further water loss once their water content decreases to approximately 1.4 g g^?1 dry weight. All larvae survive these conditions for the duration of the experiment (55 h). Eretmoptera murphyi is well adapted to survive on Signy Island, and these studies suggest that it has the ability to survive at more extreme locations at higher latitudes if it were to be inadvertently transferred to a suitable habitat.     

Morphology,egg cocoons,and transmission paths of the Antarctic leech Glyptonotobdella antarctica Sawyer and White, 1969 (Hirudinea: Rhynchobdelliformes: Piscicolidae)

The biology of piscicolids which move between different host species is poorly understood. Very little is known about leeches of Antarctic waters, though they are not uncommon in that region. This paper adds to the fundamental knowledge of the Antarctic piscicolid leech Glyptonotobdella antarctica Sawyer and White, 1969. According to the new findings the distribution of the species has to be extended to the geographical latitude of 76°08.1′S and to a depth of 665 m. Besides Glyptonotus antarcticus Eights 1853, sea urchins of the genus Sterechinus (Meissner 1900), and the benthic octopus Pareledone (charcoti?) (Joubin 1905) serve as hosts. Hitherto there were only two records on piscicolids from Octopus dofleini and none from sea urchins. Egg cocoons on the ventral side of the isopod Glyptonotus antarcticus were attributed to the leech. Most likely Glyptonotobdella antarctica moves between different hosts, i.e. Antarctic octopuses and their potential prey. Surface structures of the leech are studied by light and scanning electron microscope. The observations complement the basic knowledge on Antarctic leeches and provide arguments for phylogenetic discussions.     

Genetic diversity and evolutionary history of the Tyrrhenian treefrog Hyla sarda (Anura: Hylidae): adding pieces to the puzzle of Corsica–Sardinia biota

The Corsica–Sardinia archipelago is a hotspot of Mediterranean biodiversity. Although tempo and mode of arrival of species to this archipelago and phylogenetic relationships with continental species have been investigated in many taxa, very little is known about the current genetic structure and evolutionary history subsequent to arrival. In the present study, we investigated genetic variation within and among populations of the Tyrrhenian treefrog Hyla sarda, a species endemic to the Corsica–Sardinia microplate and the surrounding islands, by means of allozyme electrophoresis. Low genetic divergence (mean D_nei = 0.01) and no appreciable differences in the levels and distribution of genetic variability (H_E: 0.06–0.09) were observed among all but one populations (Elba). Historical demographic and isolation‐by‐distance analyses indicated that this diffused genetic homogeneity could be the result of recent demographic expansion. Along with paleoenvironmental data, such expansion could have occurred during the last glacial phase, when wide and persistent land bridges connected the main islands and a widening of lowland areas occurred. This scenario is unprecedented among Corsica–Sardinia species. Together with the lack of concordance even among the few previously studied species, this suggests either that species had largely independent responses to paleoenvironmental changes, or that most of the history of assembly of the Corsica–Sardinia biota is yet to be written. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 159–167.     

Effect of Sex and Age on the Supercooling Point of the Winter-Active Diamesa mendotae Muttkowski (Diptera: Chironomidae)

Supercooling points (SCP) of adult Diamesa mendotae Muttkowski, 1915 (Diptera: Chironomidae) were determined at 3, 5, 9, 12 and 17 days post-collection. Supercooling points were recorded using surface contact thermometry and a cooling rate of ca. 1°C min^?1. Female SCPs averaged ?22.81, ?23.76, ?23.85, ?23.65, and ?20.87°C on each date post-collection and did not differ significantly. Male SCPs were statistically similar and averaged ?21.75, ?23.53, ?23.68, ?23.66, and ?22.92°C on each date post-collection. Paired comparisons of female/male SCPs on each date post-collection did not show significant differences over time. The overall mean SCP of D. mendotae (?23.05°C) is substantially lower than values of ?5.3°C and ?5.7°C published for adults and larvae of Belgica antarctica Jacobs 1900 collected at Palmer Station (Antarctica) and ?14.2°C for larvae of Paraclunio alaskensis Coquillett 1900 collected at Vancouver Island, British Columbia. In addition, the SCP of this species appears to be lower than that of other winter-active insect species reported in the literature. Although no adults survived after the SCP was recorded, further studies are necessary to determine if D. mendotae is a freeze-intolerant insect. Nevertheless, our results suggest that a low SCP could be used as a mechanism to facilitate emergence and adult activity of this species during winter conditions.     

Description of <Emphasis Type="Italic">Gosztonyia antarctica</Emphasis>, a new genus and species of Zoarcidae (Teleostei: Perciformes) from the Antarctic Ocean

A new genus and species of zoarcid fish, Gosztonyia antarctica, is described on the basis of four specimens collected from the Bellingshausen Sea, Antarctic Ocean, at a depth of 615 m. Gosztonyia is placed in the subfamily Lycodinae and can be distinguished from all other zoarcid genera by the following combination of characters: seven branchiostegal rays, interdigitating ceratohyal–epihyal articulation, palatal arch reduced, posterior hyomandibular ramus longer than anterior, cranium narrowed, supratemporal commisure and occipital pores absent. A new species, Gosztonyia antarctica, is described and the relationships of the new genus are discussed.     

Onset of seasonal metabolic depression in the Antarctic midge Belgica antarctica appears to be independent of environmental cues

Seasonal progression is tracked in most animals by changes in daylength, thus allowing reliable synchrony with abundant food and favourable developmental conditions. In polar regions, daylength varies extensively, fluctuating at the highest latitudes from persistent light to persistent dark. The Antarctic midge Belgica antarctica has a narrow seasonal window in which to feed and develop, and previous work shows that this insect, despite having the elements of a circadian clock, remains continuously active when temperatures are permissive. The present study aims to clarify seasonal tracking in B. antarctica during the austral summer by monitoring oxygen consumption rates in a field population and in experimental groups exposed to shortened daylength, dehydration and chilling. Remarkably, during March, coordinated decreases in oxygen consumption are observed, ranging from 18% to 42%, in all treatment groups, indicating an anticipatory response to seasonal change regardless of the environmental cues. These results suggest that B. antarctica relies on an intrinsic mechanism to program metabolic depression at the onset of the long austral winter.     

Mixoscyphus antarcticus gen. nov., sp. nov. (Cnidaria, Hydrozoa, Sertulariidae), the first truly endemic genus of Antarctic benthic hydroids

A genus and species of Antarctic benthic hydroids new to science, Mixoscyphus antarcticus gen. nov., sp. nov., is described and figured. Its systematic position amongst allied genera of the family Sertulariidae is discussed. The studied material originates from the South Shetland Islands area (West Antarctica), which was collected by several Spanish and US Antarctic expeditions. Mixoscyphus gen. nov. currently represents the only complete endemic genus of Antarctic benthic hydrozoans. A discussion of other genera of benthic hydroids that are largely endemic to the Antarctic is also provided.     

Cryptic mitochondrial lineages in the widespread pycnogonid Colossendeis megalonyx Hoek, 1881 from Antarctic and Subantarctic waters

Colossendeis megalonyx Hoek, 1881 is a widespread and abundant pycnogonid in the Southern Ocean which has also been reported from the South Atlantic and South Pacific Oceans. Its strictly benthic lifestyle is expected to promote genetic differentiation among populations and ultimately facilitate speciation. On the other hand, the reported eurybathy and unknown larval stages of this species may allow Colossendeis megalonyx to maintain genetic continuity between isolated shallow-water habitats by active dispersal through the deep sea or by passive rafting on floating substrates. Thus, it remains unknown whether and to which extent geographically separated populations of Colossendeis megalonyx maintain gene flow in the Southern Ocean. We sampled 96 specimens of Colossendeis megalonyx from three stations in the Atlantic Sector of the Southern Ocean and one station from the South American continental shelf (Burdwood Bank). The genetic structure of nominal Colossendeis megalonyx as well as its phylogenetic position within the genus Colossendeis were assessed using a fragment of the cytochrome c oxidase subunit 1 gene. Our data strongly support that nominal Colossendeis megalonyx consists of at least five cryptic and one pseudocryptic mitochondrial lineages, four of which appear to be geographically restricted. Two lineages occurred at locations separated by more than 1,000 km in the Antarctic, thus indicating high levels of gene flow or recent colonization. No haplotype sharing across the Polar Frontal Zone was observed. Our results strongly suggest that cryptic speciation occurred within the genus Colossendeis. The wide biogeographic distribution range of Colossendeis megalonyx and perhaps that of other Antarctic pycnogonids should therefore be regarded with caution.     

Cold resistance in Antarctic angiosperms

Deschampsia antarctica Desv. (Poaceae) and Colobanthus quitensis (Kunth) Bartl. (Cariophyllaceae) are the only two vascular plants that have colonized the Maritime Antarctic. The primary purpose of the present work was to determine cold resistance mechanisms in these two Antarctic plants. This was achieved by comparing thermal properties of leaves and the lethal freezing temperature to 50% of the tissue (LT50). The grass D. antarctica was able to tolerate freezing to a lower temperature than C. quitensis. The main freezing resistance mechanism for C. quitensis is supercooling. Thus, the grass is mainly a freezing‐tolerant species, while C. quitensis avoids freezing. D. antarctica cold acclimated; thus, reducing its LT50. C. quitensis showed little cold‐acclimation capacity. Because day length is highly variable in the Antarctic, the effect of day length on freezing tolerance, growth, various soluble carbohydrates, starch, and proline contents in leaves of D. antarctica growing in the laboratory under cold‐acclimation conditions was studied. During the cold‐acclimation treatment, the LT50 was lowered more effectively under long day (21/3 h light/dark) and medium day (16/8) light periods than under a short day period (8/16). The longer the day length treatment, the faster the growth rate for both acclimated and non‐acclimated plants. Similarly, the longer the day treatment during cold acclimation, the higher the sucrose content (up to 7‐fold with respect to non‐acclimated control values). Oligo and polyfructans accumulated significantly during cold acclimation only with the medium day length treatment. Oligofructans accounted for more than 80% of total fructans. The degrees of polymerization were mostly between 3 and 10. C. quitensis under cold acclimation accumulated a similar amount of sucrose than D. antarctica, but no fructans were detected. The suggestion that survival of Antarctic plants in the Antarctic could be at least partially explained by accumulation of these substances is discussed.