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Chromosomal Variability in the Antarctic Insect, <I>Belgica

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Chromosomal Variability in the Antarctic Insect, <I>Belgica Chromosomal Variability in the Antarctic Insect, Belgica antarctica (Diptera: Chironomidae) WILLIAM R. ATCHLEY2 AND BRENT L. DAVIS3 ABSTRACT Ann. Entomol. Soc. Am. 72: 246-252 (1979) Chromosomal inversion polymorphism is examined in the midge Belgica antarctica Jacobs (Diptera: Chironomidae). This insect is the southernmost free-living holometabolous species and is restricted to ca. a 650-km range on western side of the Antarctic Peninsula. B. antarc- tica is a diploid species with a chromosome complement of 2n=6. Five chromosomal inver- sions were found among ca. 1200 insects from 18 localities. One inversion is sex-linked. Two inversions exhibited highly significant interpopulational differentiation in the frequency of inversion heterozygotes. Attempts to correlate inversion heterozygosity with several en- vironmental variables including ecological complexity and 3 soil variables, i.e., pH, electri- cal conductivity and percent organic material, gave non-significant results. Several hy- Downloaded from https://academic.oup.com/aesa/article/72/2/246/125422 by guest on 01 October 2021 potheses are advanced to explain the observed patterns of genetic variability. Virtually nothing is known about genetic variability area can be found in Strong (1967), Peckham (1971) and in the terrestrial insect fauna of Antarctica. The species references therein. of Antarctic insect most suitable for genetic analyses is All specimens examined in this study were mature lar- the wingless midge, Belgica antarctica Jacobs (Diptera: vae preserved in 3:1 ethyl alcohol - acetic acid fixative. Chironomidae), which is the southernmost free-living The sex of these individuals was determined by the gen- holometabolous insect. This monotypic genus is re- ital analagen method (Wiilker and Gotz 1968). Cytolog- stricted to ca. 650-km range from Elephant Island in the ical preparations were made by dissecting out the sali- northern sector of the South Shetland Islands through the vary glands into 60% proprionic acid, staining with Gerlache Straits of the Palmer Archipelago and south to lacto-proprionic-orecin and squashing in 60% proprionic 65° 27' south latitude in the Argentine Islands on the acid. western side of the Antarctic Peninsula (Wirth and Gres- sitt 1967). The mammal and bird species occurring in the Ant- Biology of Belgica arctic are considered to be either transient or marine and Preliminary studies on the biology of Belgica antarc- the terrestrial arthropod fauna includes Acarina, Collem- tica have been made by Torres (1953), Strong (1967) bola, Mallophaga, Anoplura, Siphonaptera as well as and Peckham (1971). Larval habitats include moist soil Belgica antarctica (Gressitt 1967). Since holometaboly under rocks, association with the nitrophilous alga Pra- is a derived trait in arthropods, B. antarctica would be siola, in mosses and in the rhizospheres of the grass assumed the evolutionarily most advanced free-living Deschampsia antarctica Hook. f. The larvae are not terrestrial Antarctic animal. found in association with free water. More detailed in- In this study we explore 2 themes. First, we provide formation on the larval habitats can be found in Peckham preliminary data on the chromosomal cytology of B. (1971). antarctica and explore the role of inversion heterozygos- Oviposition occurs in damp moss or similar substrate ity in adapting this species to a polar environment. Sec- with ca. 30-50 eggs laid in a gelatinous mass per ovi- ond, we examine interpopulational variation in inversion position. Hatching occurs in ca. 10 days followed by heterozygosity in a series of island populations off the dispersal of the larvae into the larval media. Individual Antarctic Peninsula. females have been observed to oviposit only once and Materials and Methods parthenogenesis is not known. Larval food appears varied consisting of dead plant A total of 1182 individuals was collected from 18 material, fungal hyphae, algae, and moss. Limited data localities on 10 small islands off the south coast of An- from Peckham (1971) indicate a preference for algae vers Island in the vicinity of Arthur Harbor (64° 46' S, over moss. 64° 02' W) on the Antarctic Peninsula. All collections Peckham (1971) has hypothesized 6 rather than the were made during the austral summer of 1974-1975 usual 4 larval instars infl. antarctica based on frequency and each sample represents a single collection. These distributions of larval head capsule length. There is also localities are described in Table 1 together with a local- some evidence that the life cycle lasts for > 1 yr in less ity code and are shown in Fig. 1. The total distance be- favorable habitats but more than one generation per year tween extreme samples (excluding EA1) was ca. 7 km. can occur in highly favorable habitats. This point re- An arrow and dollar sign denote the site of the United quires further clarification. States National Science Foundation Field Station. De- Mobility in adults is limited as a result of the loss of tailed information on the ecology of the Arthur Harbor wings and mating occurs on the substrate. Dispersal over any distance is passive involving air currents, transport 1 Received for publication Sept. 5. 1978. in the nesting materials of birds, etc. B. antarctica often 1 Dept. of Entomology. Univ. of Wisconsin. Madison 53706. 3 Dept. of Biol. Sciences. Texas Tech Univ.. Lubbock 79409. occurs in clumps of Deschampsia, moss or non-living 246 ©1979 Entomological Society of America 0013-8746/79/0202-4607$00.75/0 March 1979] ATCHLEY AND DAVIS: Belgica CHROMOSOMAL VARIABILITY 247 Downloaded from https://academic.oup.com/aesa/article/72/2/246/125422 by guest on 01 October 2021 Fie. 1.—Map of Arthur Harbor vicinity showing location of sampling sites. Insert shows the position of Anvers Island on the Antarctic Pennisula. Sample EA1 is not shown on this map. Table 1 provides an explanation of the locality codes. Table 1.—Localities, locality codes, sample sizes, frequencies of inversion heterozygotes and ecological complexity values for 18 samples of Belgica antarctica Jacobs. Total sample size and average heterozygote frequencies are given at the bottom of each column. See text for clarification of symbols. 1 2 3 4 5 6 7 8 9 10 Locality A+B+ Ecological code Locality N A B C A+B A+C B+C C '%H 1-R complexity AAI Cormorant Island 58 0.379 0.224 0.069 0.121 0.017 0.000 0.000 0.534 0.063 3 ACI Limitrophe Island 66 .439 .394 .073 .091 .030 .015 .015 .742 .083 3 AEI Hermit Island 63 .444 .492 .254 .175 .063 .063 .111 .730 .124 1 AFI Ikes Island 44 .523 .409 .136 .183 .068 .045 .000 .773 .100 3 AGI Shortcut Island 49 .571 .367 .061 .204 .020 .041 .000 .735 .091 1 AG2 Shortcut Island 77 .571 .377 .078 .195 .039 .013 .026 .727 .094 1 AJI Bonapart Point 93 .613 .247 .000 .129 .000 .000 .000 .731 .072 4 AJ3 Bonapart Point 82 .524 .317 .061 .207 .037 .000 .012 .634 .082 4 AJ4 Bonapart Point 64 .531 .438 .047 .172 .000 .000 .016 .813 .091 3 AMI Norsal Point 55 .618 .200 .200 .091 .109 .036 .018 .745 .101 1 AM5 Norsal Point 72 .542 .306 .431 .139 .139 .069 .042 .847 .139 5 AM6 Norsal Point 68 .529 .235 .000 .118 .000 .000 .000 .647 .068 4 AO1 Litchfield Island 70 .514 .443 .314 .186 .071 .086 .086 .757 .126 3 AO4 Litchfield Island 66 .530 .288 .121 .167 .061 .015 .015 .667 .089 4 AO5 Litchfield Island 62 .452 .500 .258 .161 .048 .048 .048 .855 .118 2 AO6 Litchfield Island 101 .505 .535 .020 .287 .000 .020 .000 .762 .092 3 API Strandtmann Island 43 .419 .512 .372 .140 .116 .070 .047 .884 .140 2 EAI Port Lockroy 49 .551 .592 .041 .306 .000 .000 .000 .878 .105 2 1182 0.518 0.378 0.132 0.173 0.042 0.028 0.024 0.743 0.099 248 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA [Vol. 72, no. 2 debris which serves as nest material for several species One large inversion was found in chromosome I in of Antarctic birds. this study. It occurred from just mesad of the Balbiani Cytology ring to within a few bands of the other end of the chro- mosome and therefore encompassed the nucleolar or- B. antarctica is a diploid, sexually reproducing spe- ganizing region. Only 1 individual was found to be het- cies and, like many species of chironomids, has polytene erozygous for this inversion in this study but this is salivary gland chromosomes. However, the polytene apparently the same inversion which Martin (1962) found chromosomes are much smaller and more difficult to at a frequency of ca. 8% in the single population he study than many of the commonly examined species. B. examined. The latter population occurred at a consider- antarctica has a chromosome complement of 2n=6 with able distance from those included in this study. homologues tightly synapsed. A typical salivary gland Chromosome II (Fig. 2, 3a, c) can be distinguished cell from a female individual is shown in Fig. 2. While by a medially located puff containing a nucleolar organ- the basic chromosome number in Chironominae is 2n=8 izer and be a darkly staining sub-terminal band seen at Downloaded from https://academic.oup.com/aesa/article/72/2/246/125422 by guest on 01 October 2021 (Martin et al. 1974), 2n=6 is apparently the basic chro- the bottom of Fig. 3a. For descriptive purposes, we have mosome number in the Orthocladiinae (Bauer and Beer- designated that portion of chromosome II with the sub- man 1952). terminal dark band as the left terminus and the other end Our studies have shown the occurrence of 5 chromo- as the right terminus. somal inversions in the populations examined thus far.
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