the University of Kansas Science Bulletin volume 51, no. 26. pp. 717-801 June 30, 1980 Cytotaxonomy of the Lygaeidae ( Hemipt era-Het eropt era ) b y N. Ueshima and P.D. Ashlock UNI 8121+ .•-•.•-•_••••_« :•:•:•:•:•:•:•:•:•:•: '.•.•.•.•.•.•.•.•.•••.•».•:•:•:•:• •:•:•:•:•:•-•:• '.•.•-•.•.•.•.•.•.•.•.•.•.•.•.'».•.•.••...•. THE UNIVERSITY OF KANSAS SCIENCE BULLETIN CYTOTAXONOMY OF THE LYGAEIDAE (HEMIPTERA-HETEROPTERA) 8 i I By i Norihiro Ueshima and Peter D. Ashlock 1 I II I ! g 5* 1 Vol. 51, No. 26, pp. 717-801 June 30, 1980 ANNOUNCEMENT The University of Kansas Science Bulletin (continuation of the Kansas Uni- scientific carried out at versity Quarterly) is an outlet for scholarly investigations the University of Kansas or by University faculty and students. Since its incep- tion, volumes of the Bulletin have been variously issued as single bound volumes, as two or three multi-paper parts or as series of individual papers. 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All communications regarding exchanges, sales and subscriptions should be addressed to the Exchange Librarian, University of Kansas Libraries, Lawrence, Kansas 66045. Reprints of individual papers for personal use by investigators are available should gratis for most recent and many older issues of the Bulletin. Such requests be directed to the author. The International Standard Serial Number of this pub'ication is US ISSN 0022-8850. Editor William L. Bloom Editorial Board Philip W. Hedrick Rudolf Jander Harvey Lillywhite Charles D. Michener Norman A. Slade Henry D. Stone George W. Byers, Chairman THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. 51, No. 26, pp. 717-801 June 30, 1980 Cytotaxonomy of the Lygaeidae 1 (Hemiptera - Heteroptera) NORIHIRO UESHIMA Department of Biology, Matsusaka College, Kubo-Cho, Matsusaka-shi, Mie, Japan 515 Peter D. Ashlock Department of Entomology, University of Kansas, Lawrence, Kansas 66045 CONTENTS Abstract 717 Lethaeini 757 Introduction - 717 Ozophorini 760 Materials and Methods 720 Antillocorini 760 Cytological Characteristics of the Lygaeidae 721 Targaremini _ 762 Lygaeinae 72 1 Drymini 763 Orsillinae 726 Stygnocorini 765 Metrargini 727 Cleradini 767 Nysiini 732 Myodochini 767 Orsillini 734 Udeocorini 771 Ischnorhynchinae 734 Rhyparochromini 773 Cyminae 737 Megalonotini 777 Cymini 737 Gonianotini 777 Ontiscini 739 Incertae Sedis 779 Ninini 739 Systematic and Cytological Discussion 779 Chauliopinae 739 Holokinetic chromosomes 780 Blissinae 740 Chromosome number 780 Henestarinae 746 Chromosome size 781 Geocorinae 746 The sex chromosome mechanism 783 Oxycareninae 749 The m-chromosome 785 of the sex and Pachygronthinae 75 1 Metaphase position Pachygronthini 75 1 m-chromosome 785 Teracriini 752 Species and subspecies discrimination 787 Heterogastrinae 754 Literature Cited 789 Rhyparochrominae 755 Figures Plinthisini 755 Figures 1-12. Chromosomes of Lygaeinae .... Abstract 12 subfamilies are discussed. Chromosomal complements of 330 species of Lygaeidae in 131 genera and Chromosome numbers and sizes, sex and m-chromosome characteristics including their metaphase positions, and the use of cytological data in discrimination of higher taxa, species, and subspecies are covered. No cy- of the but several taxa tological element of the Lygaeidae is unique to the family nor to any part family, may be characterized by combinations of cytological features. Introduction found. Accumulation of data has con- tinued, in Japan by Ueshima, who is re- This survey of the chromosomes of the sponsible for the cytological work and its Lygaeidae was begun early in the 1960s at the University of California, Berkeley, Contribution number 1700 from the Depart- to see information to whether pertinent ment of Entomology, University of Kansas, Law- the classification of the family might be rence, Kansas 66045. 718 The University of Kansas Science Bulletin interpretation, and in Hawaii and Kansas being the most informative and easiest to collected identified by Ashlock, who and obtain, are used in this study. Those of many of the specimens and has provided one member of each genus studied are the systematic interpretation. The work illustrated. Where significant differences has been furthered several by colleagues were found within a single genus, these who sent specimens from various parts of differences are also illustrated. Illustra- the world. tions include spermatogonial metaphase and I Work on lygaeid chromosomes was be- metaphases and II of meiosis, mostly from a view. gun early in the 1900s in the United States polar Occasionally only a lateral view is by Montgomery (190k, \90U, 1906) and shown because a suitable view was not found in the Wilson (1905a, 1905£, 1909, 1912). The polar prepara- most comprehensive contribution was that tions. Sometimes both lateral and polar views are of Pfaler-Collander (1941), who studied given, and occasionally other of are shown over 50 Finnish species of the family. stages spermatogenesis where these show Other workers in various parts of the stages significant additional information. world each have added a few to bring the total of cytologically known species to The authors would like this survey to about 75. here We add more than 250 spe- be useful to the cytologist and the He- cies to the list. The chromosomes of well miptera systematist alike. The materials over 10% of the 2,800 species listed in and methods section thus includes de- Slater's Catalogue of the Lygaeidae of the scriptions of the process of obtaining and World (1964) have now been studied. preparing chromosomes for study so that Chromosomes of Lygaeidae, like those noncytologists who are so inclined may make their of all Hemiptera (Heteroptera and Ho- own observations. Terms that are not be familiar to the moptera), holokinetic; that is, they may noncytologist have diffuse or holocentric centromeres are defined below. The actual data is pre- rather than localized centromeres as do sented by subfamilies with genera grouped most organisms. Because the centromere alphabetically after a general description of the is distributed along the length of each cytological characteristics of the tribe in the chromosome, the parts of a fragmented appropriate subfamily (or chromosome are not lost and may still large subfamily Rhyparochrominae). A of the and move to the poles at anaphase. Another summary cytological systematic unusual feature found in most Lygaeidae findings concludes the text of the paper. Tables 1 8 chromosome and in some other families of the Heterop- through compare tera is the micro- or m-chromosome. Usu- sizes of species in well-studied genera. 9 ally minute, these chromosomes are al- Table gives the modal number for each of the taxa above the level ways unpaired during meiotic prophase major genus and the of and no chiasmata are formed. During the positions the sex chromosomes first and m-chromosomes I and second anaphase, they are nega- during metaphases and II. Table 10 lists stud- tively heteropycnotic. Generally, m-chro- every species mosomes orient themselves in the center ied to date in the Lygaeidae, the source of the ring of autosomes at metaphase I of information, where the specimens were and often at metaphase II as well, with obtained, and the spermatogonial and the sex chromosomes. The cytogenetic sig- metaphase I and II chromosome comple- nificance of the m-chromosome is un- ments by number. known. Ueshima (1979) has summarized The chromosomes of spermatogenesis, heteropterous cytogenetics. His summary Cytotaxonomy of Lygaeidae 719 includes a detailed description of meiosis are not visible as discrete structures al- the sex chromosomes be dis- in Oncopeltus fasciatus (Dallas) (Lygaei- though may cernible. The diffuse is character- dae, Lygaeinae) as well as discussions of stage istic of the holocentric chromosomes, m-chromosomes, Heteroptera (Fig. 6b). the behavior and mechanism of the sex diplotene—stage during prophase in which chiasmata be evident in each chromosomes, and a list of the diploid and may of chromosomes haploid chromosome numbers of all Het- pair homologous (Fig. studied to date. As it is a 3b). teroptera such, — companion to this contribution, and should equational division the segregation pat- be consulted for further information on tern in which sister chromatids of a chromosome to hemipteran cytogenetics. segregate opposite poles (see reductional). Hemiptera have the usual stages in —chromosomes or chromo- spermatogenesis. A spermatogonial divi- heteropyenotic some regions that stain differently from sion, which is like a typical mitotic divi- the rest of the genome. Positive hetero- sion, is followed by two