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Insecta; Diptera: Chironomidae) Pacific Science (1988), vol. 42, nos. 1-2 © 1988 by the University ofHawaii Press . All rights reserved Evolutionary Relationships of the Hawaiian and North American Telmatogeton (Insecta; Diptera: Chironomidae) LESTER J. NEWMAN! ABSTRACT: Species of Telmatogeton and the closely related genus Para­ clunio generally live on the rocky shores of the intertidal zone. Species of Tel­ matogeton have evolved from the marine environment into torrential freshwater streams of the Hawaiian Islands . An analysis of the banding sequences of the polytene chromosomes of species of Telmatogeton and Paraclunio from the Hawaiian Islands and North America suggests that there were at least two separate invasions from the marine to freshwater environments. One is through the marine species T. japonicus to an undescribed freshwater species found on east Maui . The other invasion requires a marine hypothetical species that gave rise to the Hawaiian freshwater species T. abnormis and T. torrenticola. SPECIES OF THE FAMILY Chironomidae are ical feature distinguishing the genera is a generally found in aquatic or semiaquatic sexual dimorphism of the front femur and environments. Species in only 13 genera of tibia . These structures are narrow and undif­ chironomids, including the Telmatogeton ferentiated in males and females of Telmato­ and Paraclunio, occupy the marine environ­ geton and in females of Paraclunio. The front ment (Hashimoto 1976, Sublette and Wirth femur of Paraclunio males is broad, and an 1980). The life cycle of most species of Tel­ angular projection of the distal portion of matogeton and Paraclunio is spent on the the femur fits into a corresponding notch in rocky shores of the intertidal zone. In the the base of the tibia. Because of the morpho­ Hawaiian Islands, species of Telmatogeton logical similarity of the two genera, the genus have evolved from the marine environment Paraclunio should be synonymized with the into torrential freshwater streams (Wirth Telmatogeton (Peter Cranston, personal 1947). communication). The Telmatogeton and Para­ Wirth (1947, 1959) describes the discovery clunio are thus essentially treated together in and naming of species of the Telmatogeton this discussion. and Paraclunio . The genus Telmatogeton was The natural history and ecology of the established by J. R. Schiner in 1866, and the Telmatogeton and Paraclunio are described first species, T. sancti-pauli, was described by Kronberg (1986), Morley and Ring by him in 1868 from specimens collected on (1972), and Robles (1982). Two important St. Paul Island in the Indian Ocean. D. W. points relative to the current discussion are Coquillett described the second species, features that lead to the isolation of popula­ T. alaskensis, from specimens collected from tions and the adaptation of species to a Yakutat, Alaska, during the Harriman Ex­ severe environment. Adults of both marine pedition of 1899. The first Hawaiian fresh­ and freshwater species exhibit a peculiar water species was described by F. W. Terry swarming behavior; they are generally seen in 1913. J. J. Kieffer established the marine running about on intertidal rocks and on genus Paraclunio in 1911 for P. trilobatus, stream banks seeking mates. Adults are not collected on the coast of California, and in strong fliers and thus may have limited 1915, J. R. Malloch moved T. alaskensis to vagility. Adults of the marine species are be­ the genus Paraclunio. The major morpholog- lieved to be short-lived ; they emerge on an outgoing tide, mate and lay eggs, and are 1 Portland State University, Department of Biology, washed out to sea on the incoming tide. Eggs P.O. Box 751, Portland, Oregon 97207. of the marine species are laid singly in rock 56 Evolutionary Relationships of Telmat ogeton-r-t-aswws i« 57 crevices, and larvae feed by grazing on the cannot be described in terms of simple in­ , thin film of algae growing on the rock sur­ versions. These are referred to as complex faces and on macroscopic green and red portions of an arm or complex arms. algae. Larvae build silky tubes from their Chromosome numbers in the Telmato­ salivary gland secretion; pupation occurs in geton and Paraclunio range from n = 7 to reinforced tubes. The tubes protect larvae n = 3/4 (Newman 1977). It has been pro­ and pupae of the marine species from wave posed that the primitive number is n = 7 and action at high tide and predation and des­ that the other numbers are derived by cen­ iccation at low tide, and thus allow larvae tric fusions. Each centric fusion reduces the and pupae of the freshwater species to live chromosome number by 1. Telmatogeton in torrential streams and waterfalls in the hirtus (n = 3/4) has an XY1 Y z sex chromo­ Hawaiian Islands . some system that may be seen in both polytene In 1971, Hampton Carson discovered the and nonpolytene chromosomes. Telmatogeton Hawaiian Telmatogeton as a material suit­ abnormis (n = 4) has an XY chromosome able for chromosome analysis (personal system that may be seen only in the polytene communication). He found the giant poly­ chromosomes. Other species do not have dif­ tene chromosomes of the larval salivary ferentiated sex chromosomes. glands to be well banded and likely to give Inversion sequences have been used to information relative to the evolution of the gain insight into possible evolutionary rela­ freshwater species. This proved to be true, tionships among species. Species that share for Newman (1977) established a standard inversions are more closely related than band map and proposed a pro visional set of species that do not share inversions. Carson evolutionary relationships for the Hawaiian (reviewed in Carson and Yoon 1982) used freshwater Telmatogeton. this type of analysis to determine the evolu­ All the species examined to date have tionary relationships of over 100 species of polytene chromosomes suitable for detailed Hawaiian Drosophila. Similar techniques are band analysis. Each species has six long used here to propose a set of relationships in chromosome ' arms with well-developed which there were at least two separate in­ bands and a single unbanded short arm . The vasions of marine Telmatogeton into fresh six long arms were designated by the letters water in the Hawaiian Islands . A-F and the short arm by the letter G. A sequence of bands within arms A-F com­ mon to most of the Hawaiian freshwater species was chosen as the Telmatogeton MATERIALS AND METHODS standard (Newman 1977). The chromosome The giant banded polytene chromosomes arms of most of the species examined have were examined from cells of the salivary the standard sequence or differ from it by gland of fourth instar larvae fixed in acetic paracentric inversions. A paracentric inver­ alcohol and stained in lactoacetic orcein. sion is a chromosomal mutation in which a Nonpolytene chromosomes for the determi­ series of bands that does not contain the cen­ nation of chromosome numbers were derived 0 tromere is rotated 180 • The specific break from lactoacetic orcein squashes of testes points of the inversion may be determined by and other developing larval tissues (details in comparing the inverted and standard band Newman 1977). sequences. Inversions are designated by giv­ ing the arm letter of the inversion with a sub­ script arabic number. An inversion may be homozygous in all animals of a population RESULTS or species; that is, it is fixed or it may exist A list of the described species of Telmato­ along with the standard as a polymorphism. geton and Paraclunio is given in Table 1. In some cases, the rearrangement of bands in Band sequence and chromosome number a portion or even an entire arm may be so data for all the species examined to date are complicated by multiple inversions that it given in Table 2. Included are data for an un- 58 PACIFIC SCIENCE, Volume 42, January/Jul y 1988 TABLE I S PECIES OF Te/matogeton AND Parac/unio SPECIES GEOGRAPHIC DISTRIBUTION Te/matogeton, marine T.japonieus Tokunaga Japan, Hawaii,* Florida, Australia, Baltic Sea* T. paeifieus Tokunaga Japan, Hawaii* T. pusillum Edward s Marquesas, Mariana, Micronesia T. maeswaini Wirth California* T. /atipennis Wirth Revillagigedo Island s (Pacific Ocean, Mexico) T. sancti-pauli Schiner South Africa, St. Paul Island (Indian Ocean) T. minor Kieffer South Africa T. australieus Womersley Australia T. antipodensis Sublette and Wirth Antipodes Islands (New Zealand) T. mortoni Leader New Zealand T. simplicipes Edwards South America (Ancud, Chile) T. troeanteratum Edwards South America (Ancud, Chile) T. at/antieum Oliveira South America T. nanum Oliveira South America Te/matogeton , Hawaiian freshwater T. hirtus Wirth Kauai* T. abnormis Terry Kauai,*Oahu Ti fluviatilis Wirth Oahu * T. williamsi Wirth Oahu T. torrentico/a Terry Molokai,* Maui,* Hawaii* Parac/unio, marine P. tri/obatus Kieffer Washington ,* Oregon,* California* P. spinosus Hashimoto California* P. a/askensis Coquillett Alaska,* British Columbia,* Washington,* Oregon,* California* List ofspecies from: Wirth (1947,1959), Hashimoto (1976), Sublette and Wirth (1980). Species moved into the genus Telmatogeton by Sublette and Wirth (1980) are not included in this table. •Chro moso mes examined. described species from east Maui designated geton japonicus, T. macswaini, and T. abnor­ as T. new species-I (T. n. sp.-I). Most of the mis are fixed for two inversions (C3,4 ) in species were discussed in Newman (1977); which inversion C4 is included within the new data are reported here for T. japonicus, limits of C 3 (Figure I). Two of the four break T. macswaini, the three Paraclunio species, points appear to be at the same site in the and an unidentified Telmatogeton species polytene chromosome. Arm C of T. n. sp.-l from southeast Maui. Adults, required for is complex but has one of the C3 break points. species identification, were not collected for Telmatogeton japonicus and T. n. sp.-I share larvae of the southeast Maui population.
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