A peer-reviewed open-access journal ZooKeysCytogenetics 154: 31–70 (2011) of the true bug infraorder (, ): a review 31 doi: 10.3897/zookeys.154.1953 Research article www.zookeys.org Launched to accelerate biodiversity research

Cytogenetics of the true bug infraorder Cimicomorpha (Hemiptera, Heteroptera): a review

Valentina G. Kuznetsova1, Snejana M. Grozeva2, Seppo Nokkala3, Christina Nokkala3

1 Zoological Institute RAS, Universitetskaya emb. 1, St Petersburg 199034, Russia 2 Institute of Biodiversity and Ecosystem research, BAS, Tsar Osvoboditel blvd, 1, Sofia 1000, Bulgaria3 Laboratory of Genetics, De- partment of Biology, University of Turku, 20500 Turku, Finland

Corresponding author: Valentina G. Kuznetsova ([email protected])

Academic editor: Pavel Štys | Received 23 August 2011 | Accepted 5 December 2011 | Published 12 December 2011

Citation: Kuznetsova VG, Grozeva SM, Nokkala S, Nokkala C (2011) Cytogenetics of the true bug infraorder Cimicomorpha (Hemiptera, Heteroptera): a review. ZooKeys 154: 31–70. doi: 10.3897/zookeys.154.1953

Abstract The Cimicomorpha is one of the largest and highly diversified infraorders of the Heteroptera. This group is also highly diversified cytogenetically and demonstrates a number of unusual cytogenetic characters such as holokinetic chromosomes; m-chromosomes; multiple sex chromosome systems; post-reduction of sex chromosomes in meiosis; variation in the presence/absence of chiasmata in spermatogenesis; different types of achiasmate meiosis. We present here a review of essential cytogenetic characters of the Cimico- morpha and outline the chief objectives and goals of future investigations in the field.

Keywords Hemiptera, Heteroptera, Cimicomorpha, holokinetic chromosomes, telomeres, NOR, chromosome number, m-chromosomes, sex chromosomes, B-chromosomes, meiosis

Introduction

The Heteroptera, or true bugs, are a diversified group of displaying a number of unusual and sometimes unique cytogenetic characters such as holokinetic chromo- somes, m-chromosomes, multiple sex chromosome systems, sex chromosome post- reduction and occasionally pre-reduction in male meiosis, variation in the presence/ab- sence of chiasmata in spermatogenesis, different types of achiasmate meiosis and oth-

Copyright V. G. Kuznetsova et al. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 32 V. G. Kuznetsova et al. / ZooKeys 154: 31–70 (2011) ers. The pioneer investigators of true bug cytogenetics were Henking (1891), McClung (1902) and Wilson (1905, 1909). It should be noticed that Hermann Henking (1891) and his object, the firebug Pyrrhocoris apterus Linnaeus, 1758 (: Pyr- rhocoridae), deserve the credit for the discovery of a relation between chromosomes and sex determination in . Since that time chromosomal sex determination has become more and more widely accepted among biologists. The cytogenetics of the Heteroptera has been firstly comprehensively reviewed by Ueshima (1979) and shortly afterwards by Manna (1984). Ueshima’s (1979) superior monograph covers characteristics of all but one (Enicocephalomorpha, for which in- formation is lacking to this day) heteropteran infraorders. However, the infraorders are cytogenetically unequally explored. Since Ueshima’s publication a large body of new cytogenetic data on the Het- eroptera has been obtained, including those on the cimicomorphan families Tin- gidae (Nokkala and Nokkala 1984a, Grozeva and Nokkala 2001), s.str. (Nokkala and Nokkala 1986a, Wang et al. 2003), (Nokkala and Grozeva 2000), (Grozeva and Nokkala 2002, Poggio et al. 2009, Grozeva et al. 2010, 2011), (Pérez et al. 2004, Severi-Aguiar et al. 2006, Poggio et al. 2007, 2011, Panzera et al. 2010, Bardella et al. 2010 ), s.str. (Nokkala and Nokkala 1984b, Kuznetsova and Maryańska-Nadahowska 2000, Kuznetsova et al. 2004, 2007, Kuznetsova and Grozeva 2008, Angus et al. 2008), and (Nok- kala 1986a, Nokkala and Nokkala 1986b, Grozeva et al. 2006, 2007, 2011, Grozeva and Simov 2008a, b, Grozeva and Simov 2009). At present, the families Miridae and Reduviidae (data are available for 196 in 83 genera and for 148 species in 45 genera, respectively), and the monospecific family Joppeicidae are the most extensively studied, whereas the families Anthocoridae s.str. (5 species, 3 genera), (3 species, 2 genera), and Microphysidae (2 species, 2 genera) are the least studied. In the three remaining families, data are available for 53 species (20 genera) in Cimicidae; 29 species (7 genera) in Nabidae s.str.; and 28 species (17 genera) in (Ta- ble 1). At present, no cytogenetic data are available for the families Pachynomidae, Vianaididae (often included in the Tingidae), Velocipedidae and Medocostidae (both sometimes included in the Nabidae s.l.), (possibly partly belonging to the Pentatomomorpha), Plokiophilidae, and and (prior to Schuh and Štys (1991), classified within Anthocoridae s.l.). The Cimicomorpha is one of the largest and highly diversified heteropteran in- fraorders. Although this group has attracted considerable interest for several reasons (disease transmission in the , evolution of host-plant relationships in the Miridae, maternal care in the Tingidae and so on; Schuh et al. 2009), cimicomorphan higher-level relationships are complex both at the family and tribal levels and subjected to several recent analyses (Schuh and Štys 1991, Schuh 1995, Schuh et al. 2009). Cy- togenetically considered, Cimicomorpha appear likewise sufficiently heterogeneous. The aim of the present paper is to synthesize main data available concerning cytoge- netic characteristics of cimicomorphan true bugs and to gain a better insight into the cytogenetic evolution within different families and the Cimicomorpha as a whole. A Cytogenetics of the true bug infraorder Cimicomorpha (Hemiptera, Heteroptera): a review 33 further aim is to outline the chief objectives and goals of future investigations in the field. The principle cytogenetic features of Cimicomorpha are summarized in Table 1 and in Figures 1 and 2.

100

90

80

70 Microphysidae 60 Joppeicidae Anthocoridae 50 Polyctenidae Cimicidae 40 Tingidae Nabidae 30 Miridae Reduviidae 20

10

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 78 Figure 1. Autosome numbers’ range in Cimicomorpha. X-axis denotes the diploid number of auto- somes, Y-axis shows the number of species

200

180

160

140 Polyctenidae Microphysidae

120 Joppeicidae Anthocoridae 100 Nabidae Cimicidae 80 Tingidae Reduviidae 60 Miridae

40

20

0 X Y X Y 2 X Y 3 X Y 4 X Y 5 X Y n XO X2O Figure 2. Distribution of sex chromosome systems in Cimicomorpha. Different sex chromosome sys- tems are plotted on the X-axis. Y-axis shows the number of species. Xn - the number of X-chromosomes exceeds 5. 34 V. G. Kuznetsova et al. / ZooKeys 154: 31–70 (2011) References (Linnaeus, 1763) (Montgomery A. cristatus (Linnaeus, 1763) (Montgomery 1944: as Troedsson 1909, 1901a, Payne 1886) Uhler, Prionidus Payne 1909, Montgomery 1901a 1909, Montgomery Payne Ueshima 1979 Payne 1912, Ueshima 1979, Poggio et al. 2007 1912, Ueshima 1979, Poggio Payne Manna 1951, Manna and Deb-Mallick 1981 and Deb-Mallick 1951, Manna Manna Ueshima 1979 Ueshima 1963b Ueshima 1963b Y (1) 1944 Troedsson 1909, 1910, Payne 5 X 4 X Y Y et al. 2007 Poggio Y 1959b 1958, Jande 1951, Banerjee Manna 3 3 3 3 2n X X X Y Y Y X 1988 Wangdi and Dey 1959a Jande 1909 Payne 2 2 2 2 2 2 2 X X X X X X X 1 1 1 1 1 1 1 (number of species) n=16 (1) 28+X0 18+XY (number of species studied) Stål, 1860 (1) Stål, Bactrodes 24+XY Stål, 1866 (1) Stål, Cydnocoris 24+X Stål, 1866 (2) Stål, 24+X Hahn, 1831 (5) Hahn, Apiomeris 22+XY Curtis, 1833 (1) Curtis, Coranaus 1859 (1) Stål, Laporte, 1833 (2)Harpactor 24+X 24+X 24+X Burmeister, 1835 Ectrychotes Burmeister, (2) Hahn, 1831 (1) Hahn, Arilus 22+X Bergroth, 1903 (1) Bergroth, 32+XY Stål, 1866 (1) Stål, Wolf, 1881 (1) Wolf, 14+XY Illiger, 1807 (2) Illiger, 26+2m+XY et al. 2011 1957, Poggio Piza Stål, 1862 (2) Stål, 20+X Taxa Subfamily Bactrodinae Stål, 1866 Stål, Bactrodinae (1/1) Ectrichodiinae Amyot and Amyot Ectrichodiinae Serville, 1843 (1/2) Emesinae Amyot and Amyot Emesinae Serville, 1843 (3/3) Stål, 1859 Stål, Hammacerinae (1/2) Amyot and Amyot Harpactorinae Serville, 1843 (18/35) Family ab Reduvioidea Superfamily Latreille, Reduviidae 1807 (45/148) (genera/species studied) T Schuh (1995) (2008) for the Reduviidae; Weirauch (1986–1989) and and Putshkov (1990), Putshkov Maldonado generally follows systematics at subfamily level Štys (1991); the systems in Cimicomorpha (The systematics at superfamily is after Schuh and numbers and sex chromosome and family level le 1. Chromosome of families) (1995) for the rest (1991) and Schuh Slater Schuh and Štys s.str.; Kerzhner (1996) for the Nabidae and Schuh (2011) for the Miridae; Cytogenetics of the true bug infraorder Cimicomorpha (Hemiptera, Heteroptera): a review 35 References Dey and Wangdi 1988, Satapathy and Patnaik and Patnaik 1988, Satapathy Wangdi and Dey 1989 Montgomery 1901a*, Payne 1909, 1912, 1901a*, Payne Montgomery 1951 1944, Manna Troedsson Jande 1959a Jande Ueshima 1979 Stål, 1862 (Payne 1909: as 1862 (Payne exsanguis Stål, 1900) Diplacodus Kirkaldy, et al. 2007 Poggio Stål, 1859 (Toshioka 1936, 1859 (Toshioka armatissimus Stål, P. 1958) Banerjee armatissimus (Jande 1960) P. 1959b Jande Jande 1959b Jande Y (1) 5 X 4 Y Y 1916 1912, Goldsmith Payne 1959b, 1958, Jande 1951, Banerjee Manna Y (5) X Y Y 1959a 1951, Jande Manna 1947 Yoshida 1933, Toshioka 3 3 3 3 3 3 2n Y 1989 and Patnaik Satapathy X X Y X X X 1909 Payne X Y (2) Y Ueshima 1979 2 2 2 2 2 2 2 2 2 2 X X X X X X X X X X 1 1 1 1 1 1 1 1 1 1 (number of species) 20+X 24+XY 24+X 20+X 10+XY 12+XY 24+X 20+X Genus (number of species studied) Spinola, 1840 (1) Spinola, Heniartes 22+XY Hahn, 1834 (3)Rhynocoris Hahn, 24+X 1865 (1) Stål, Velinus 1802 (2) Fabricius, Zelus 24+X 24+XY Stål, 1859 (1) Stål, 24+X Amyot and Serville, Amyot Sycanus 1843 (2) Rasahus and Serville, Amyot 1843 (1) Stål, 1858 (2) Stål, Polididus 10+XY (1) 1981 and Deb-Mallick Manna and Serville, Amyot 1843 Sinea (6) 1837 (1) Spinola, 26+XY Lophocephala Laporte, 1833 (1) 24+X Stål, 1863 (1) Stål, Androclus 20+XY Bergroth, 1905 (1) Bergroth, 24+X Mayr, 1865 (3) Mayr, Ectomocoris 20+XY (1) Taxa Subfamily Stål, 1859 (4/6) Stål, Peiratinae Family 36 V. G. Kuznetsova et al. / ZooKeys 154: 31–70 (2011) References Jande 1959a, Ueshima 1979, Manna and Deb- 1959a, Ueshima 1979, Manna Jande 1989 and Patnaik 1981, Satapathy Mallick Banerjee 1958 Banerjee 1959a Jande et al. 1972, Panzera Ueshima 1966a, Schreiber et al. 2010 1835) (Schreiber megistus (Burmeister, P. 1956: as Mestor 1950, Barth and Pellegrino 1904) Kirkaldi, Ueshima 1979 Ueshima 1966a Montgomery 1901a* Montgomery Payne 1912, Ueshima 1979 Payne Ueshima 1979 Jande 1959a Jande Y 1959b Jande Y 1912 Payne Y et al. 2007 Poggio 4 4 4 X Y (1) Y (2) X X 3 3 3 3 3 2n X Y (4) X X X Y (1) X Y Y Y et al. 2010 Panzera Y (1) et al. 2010 Panzera Y (7) 2010 et al. 2004, Frìas-Lasserre Pérez Y (1) 2 2 2 2 2 2 2 2 2 2 2 2 2 X X X X X X X X X X X X X 1 1 1 1 1 1 1 1 1 1 1 1 1 (number of species) 22+X 20+X 20+X 18+X 26+XY 20+XY 20+X 26+XY (1) 22+X Usinger, 1939 (1) Usinger, Genus (number of species studied) Swederus, 1787 Swederus, (1) Stål, 1859 (1) Stål, 20+X Stenopoda Laporte, 1833 (1)Stenopoda 20+X 1938 (1) Barber, 20+XY (1) Ueshima 1966a and Gajardo Mazza, Mepraia 1940 (3) Jörg, 1879 (9) Berg, Panstrongylus 18+X Latreille, 1802 (1) Latreille, Phymata 26+XY Fabricius, 1775 (2) Fabricius, 20+XY (1) 1817 (2) Germar, Pygolampis 22+XY (1) 1859 (2) Stäl, Belminus 20+X Stäl, 1859 (2) Stäl, 20+X Reuter, 1881 (1) Reuter, Pasiropsis 24+X Kirkaldi, 1900 (1) Kirkaldi, Staliastes 24+XY Klug, 1830 (7)Oncocephalus 20+X Taxa Subfamily Phymatinae Laporte,Phymatinae 1832 (2/2) Jeannel, 1919 Jeannel, Triatominae (8/84) Reduviinae (3/4) Reduviinae Stenopodinae Amyot and Amyot Stenopodinae Servielle, 1843 (4/11) Family Cytogenetics of the true bug infraorder Cimicomorpha (Hemiptera, Heteroptera): a review 37 References Schreiber and Pellegrino 1950, Barth 1956, 1950, Barth and Pellegrino Schreiber et et al. 2006; Panzera Ueshima 1966a, Bargues al. 1995, 2004, 2006, 2010 (Payne 1909, sanguisuga (LeConte, 1855) (Payne T. Laporte, et al. 2010: as Conorhinus Panzera 1832) 1773) (Manna Geer, rubrofasciatus (De T. ) et al. 2010: as Conorhinus 1950, 1951, Panzera (Bergroth 1911) (Schreiber and 1911) (Schreiber (Bergroth R. coreodes Bergroth, 1950: as Pellegrino 1956, Ueshima 1966a, Petitpierre 1911), Barth et al. 2010 1996, Panzera Nokkala and Grozeva 2000 and Grozeva Nokkala Curtis, 1833 (Grozeva, 1833 (Grozeva, L. pselapfiformis Curtis, unpublished) Ueshima 1979 Kumar 1971 Kumar Grozeva and Simov 2008b and Simov Grozeva Y (2) 3 2n Y (1) Y (21) X Y Y 2 2 2 2 2 X X X X X 1 1 1 1 1 (number of species) 20+X 20+XY (25) 20+X 20+X 22+X 12+XY 32+XY Genus (number of species studied) Stål, 1859 (15) Stål, 20+XY Bärensprung, Bärensprung, 1857 (1) Laporte, 1832 (51)Triatoma 18+X Curtis, 1833 (1) Curtis, 12+XY Putton, 1881 (1) Putton, Joppeicus 22+XY Schumacher, 1917 Bryocoropsis Schumacher, (1) Fallén, 1829 (1)Bryocoris Fallén, 32+XY Taxa Subfamily 1860 Baerensprung, (9/27) Family Superfamily Microphysoidea Superfamily Dohrn, Microphysidae 1859 (2/2) Superfamily Joppeicoidea Superfamily 1910 Reuter, Joppeicidae (1/1) Superfamily Superfamily 1833 Hahn, Miridae (83/196) 38 V. G. Kuznetsova et al. / ZooKeys 154: 31–70 (2011) References Grozeva et al. 2006, 2007 Grozeva Slack 1938b*, Leston 1957, Grozeva 2003, 1938b*, Leston 1957, Grozeva Slack 2008b and Simov Grozeva Slack 1938b, Leston 1957, Kumar 1971, Leston 1957, Kumar 1938b, Slack and Simov Akingbohungbe 1974*, Grozeva 2008b Kumar 1971 Kumar Kumar 1971 Kumar 1971 Kumar Southwood and Leston 1959*, Takenouchi Takenouchi and Leston 1959*, Southwood 1968, Akingbohungbe 1974*, and Muramoto et al. 2011 Grozeva D. lutescens Schilling, 1837 (Leston 1957: as (Schilling, 1837)) Camptobrochis Akingbohungbe 1974* Takenouchi and Muramoto 1967, 1972a, b, 1967, 1972a, b, and Muramoto Takenouchi 1973a, Akingbohungbe 1974* Muramoto Y (1) 3 X 2 Y (1) Y (1) X 3 3 1 2n Y (1) Y (1) X X 2 2 2 2 X X X X 1 1 1 1 (number of species) 26+XY (1) 46+X 46+XY (8) 46+2m+X 46+X 40+XY (1) 44+XY(1) 44+X 18+XY (1) 24+X 32+XY (11) 34+XY (1) 30+2m+XY (2) 32+XY 34+XY (1) 22+XY (2) 24+XY (1) 26+XY (1) Genus (number of species studied) Fieber, 1858 (1) Fieber, Campyloneura 32+XX ( ♀♀ ) 2008b and Simov Grozeva Dahlbom, 1851(2) Dahlbom, Molanocoris 32+XY Distantiella China, 1944 (1) 26+XY Fieber, 1858 (15)Dicyphus Fieber, 36+XY (1) Haglund, 1895 (1) Haglund, Sahlbergella 26+XY Signoret, 1858 (2) Signoret, Helopeltis 16+XY (1) Fieber, 1858 (3) Fieber, Macrolophus 24+XY (1) Kirschbaum, 1856 Deraeocoris (15) Reuter, 1876 (2) Reuter, 32+XY (1) Reuter, 1896 (7) Reuter, 20+XY (1) Taxa Subfamily Deraeocorinae Douglas Douglas Deraeocorinae and Scott, 1865 (2/17) Hahn, 1833 Hahn, Mirinae (34/73) Family Cytogenetics of the true bug infraorder Cimicomorpha (Hemiptera, Heteroptera): a review 39 References A. lineolatus (Goeze, 1778) (Schachow 1932*) 1778) (Schachow A. lineolatus (Goeze, 1941: as / A. lineolatus (Ekblom 1881, Leston 1957, chenopodii /Westhoff, Akingbohungbe 1974*) 1832) (Akingbohungbe 1974*: (Say, A. rapidus 1832) Say, rapidus as Calocoris (Mongomery 1901a, 1906: as (Mongomery A. rapidus ) rapidus Calocoris Leston 1957, Muramoto 1973a Leston 1957, Muramoto Southwood and Leston 1959* Southwood (Slack 1938b*, C. ater (Linnaeus, 1758) (Slack Akingbohungbe 1974*) 1986b) and Nokkala C. ater (Nokkala Muramoto 1973a Muramoto De Geer, 1773 (Slack 1773 (Slack Geer, De C. fulvomaculatus 1858) Fieber, 1938b*: as Calocoris C. norvegicus1938b*: as Gmelin, 1790 Slack ) Calocoris C. norvegicus) (Leston 1957: as Calocoris Akingbohungbe 1974* Takenouchi and Muramoto 1970 and Muramoto Takenouchi Reuter, 1896 (Takenouchi and 1896 (Takenouchi C. saundersi Reuter, 1967: as Orthops ) Muramoto Takenouchi and Muramoto 1968 and Muramoto Takenouchi Akingbohungbe 1974* 2n 0 2 X 1 (number of species) 12+2m+XO 30+XY 2n=28 26+X 30+2m+XY 30+XY 32+XY 28+2m+XY (1) 2003 Grozeva Genus (number of species studied) Apolygus China, 1941 (3)Apolygus 32+XY Reuter, 1896 (1) Reuter, Camptozygum 32+XY Fabricius, 1803 (1) Fabricius, 32+XY Fieber, 1858 (1) Fieber, Charagochilus 32+XY Fieber, 1858 (2) Fieber, 32+XY (1) Provancher, 1872 (1)Collaria Provancher, 12+XY Distant, 1883 (1) Distant, Creontiades 30+XY Reuter, 1903 (1) Reuter, Cyphodemidea 32+XY Fieber, 1858 (2) Fieber, Dichrooscytus 32+XY (1) Akingbohungbe 1974* Stål, 1871 (1) Stål, Eurystylus 30+XY Stål, 1862 (1) Stål, Garganus 32+XY Taxa Subfamily Family 40 V. G. Kuznetsova et al. / ZooKeys 154: 31–70 (2011) References (Fabricius, 1777) (Slack 1938b*: 1777) (Slack G. sexguttatus (Fabricius, ) as Calocoris Montgomery 1901a, 1906, Akingbohungbe Montgomery 1974* Montgomery 1901a, Slack 1938b*, Leston 1901a, Slack Montgomery 1957, Akingbohungbe 1974* (Fabricius, 1781) (Leston 1957: L. tripustulatus (Fabricius, ) as Lygus Akingbohungbe 1974* and Leston 1959*, Muramoto 1973a, and Leston 1959*, Muramoto Akingbohungbe 1974* and 1876 (Takenouchi L. rubripes Jakovlev, ) 1970: as Adelphocoris Muramoto Akingbohungbe 1974*, Ueshima 1979 (Linnaeus, 1758) (Montgomery L. pratensis 1901a, 1906) 1932*) (Schachow L. pratensis 1939*) (Geitler L. pratensis Leston 1957 (Geoffroy, 1785) (Leston 1957) (Geoffroy, M . recticornis et al. 2011) (Grozeva M . recticornis Akingbohungbe 1974* Takenouchi and Muramoto 1968 and Muramoto Takenouchi (Linnaeus, 1758) (Muramoto O. campestris (Linnaeus, 1758) (Muramoto 1973a) 2n (number of species) 30+XY 32+XY n=19? 30+2m+X0 32+XY 30+XY Genus (number of species studied) Douglas and Scott, Douglas 1868 (1) Distant, 1884 (1) Distant, Horcias 32+XY Fieber, 1858 (1)Leptopterna Fieber, 32+XY Fieber, 1858 (1) Fieber, 32+XY Slater, 1956 (1) Slater, Litomiris 32+XY Reuter, 1875 (11) Reuter, Lygocoris 32+XY (10) 1939*, Leston 1957, Southwood Geitler Hahn, 1833 (5) Hahn, Lygus 32+XY (4) and Leston 1959*, Leston 1957, Southwood Fieber, 1858 (1) Fieber, Megacoelum 30+XY Fieber, 1858 (1) Fieber, Megaloceroea 32+XY Reuter, 1876 (3) Reuter, Neurocolpus 32+XY Fieber, 1858 (1) Fieber, 12+2m+X0 1932 Schachow Distant, 1904 (1) Distant, Onomaus 30+XY Fieber, 1858 (1)Orthops Fieber, 22+XY Taxa Subfamily Family Cytogenetics of the true bug infraorder Cimicomorpha (Hemiptera, Heteroptera): a review 41 References O. campestris (Akingbohungbe 1974*) Slack 1938b, Leston 1957, Southwood and Leston 1957, Southwood 1938b, Slack Leston 1959*, Akingbohungbe 1974* (De Geer, 1773) Geer, (De D. flavoquadrimaculatus 2011) and Simov (Grozeva Montgomery 1901a, Montgomery Akingbohungbe 1974* Leston 1957, Akingbohungbe 1974* Calocoris 1986b: as Calocoris and Nokkala Nokkala quadripunctatus (Villers, 1789) Leston 1957 1938b*, Leston 1957 Slack (Fabricius 1794) (Slack 1938b*, 1794) (Slack S. binotatus (Fabricius 1967) and Muramoto Takenouchi S. binotatus (Leston 1957) S. binotatus (Akingbohungbe 1974*) Akingbohungbe 1974* Takenouchi and Muramoto 1967, and Muramoto Takenouchi Akingbohungbe 1974* Akingbohungbe 1974* Leston 1957 Akingbohungbe 1974* Y and Leston 1959* Southwood 2n 2 X 1 (number of species) 32+XY 30+XY (1) 32+XY 32+XY (1) 30+XY 32+XY 2n=32-34 32+XY 22+XY 22+XY (3) Genus (number of species studied) Fallén, 1814 (9) Fallén, Phytocoris 32+XY (8) Reuter, 1875 (2)Dryophilocoris Reuter, 32+XY (1) 2011 and Simov Grozeva Reuter, 1876 (1) Reuter, Poecilocapsus 32+XY Hahn, 1831 (3) Hahn, Polymerus 32+XY Wagner, 1968Rhabdomiris Wagner, 30+XY Stenodema Laporte, 1833 (2)Stenodema 30+XY (1) Jakovlev, 1877 (2) Jakovlev, Stenotus 40+XY (1) 1967 and Muramoto Takenouchi Distant, 1883 (1) Distant, Taedia Fieber, 1858 (2) Fieber, Trigonotylus 32+XY Uhler, 1878 (1) Uhler, Tropidosteptes 32+XY Kolenati, Kolenati, Brepharidopterus 1845 (1) Reuter, 1876 (4) Reuter, 18+XY (1) Hahn, 1834 (1) Hahn, Cyllecoris 21+ X Taxa Subfamily Van Duzee, Duzee, Van 1916 (16/33) Family 42 V. G. Kuznetsova et al. / ZooKeys 154: 31–70 (2011) References (Leston 1957) D. flavoquadrimaculatus Akingbohungbe 1974* Southwood and Leston 1959* Southwood Akingbohungbe 1974* Akingbohungbe 1974* Akingbohungbe 1974* Takenouchi and Muramoto 1967 and Muramoto Takenouchi Muramoto 1972b, and Muramoto Takenouchi 1974 1973b, Slack 1938b*, Leston 1957, Southwood and 1938b*, Leston 1957, Southwood Slack Leston 1959*, Akingbohungbe 1974* 1841) (Leston (C Sahlberg, O. flavosparsus 1957) (Akingbohungbe 1974*) O. flavosparsus Akingbohungbe 1974* Akingbohungbe 1974* Akingbohungbe 1974* Akingbohungbe 1974* Southwood and Leston 1959*, Akingbohungbe Southwood 1974* Southwood and Leston 1959* Southwood Akingbohungbe 1974* Slack 1938b*, Leston 1957, Southwood and 1938b*, Leston 1957, Southwood Slack Leston 1959*, Akingbohungbe 1974* 2n Y 2 X 1 (number of species) 34+X 32+XY 28+XY 24+XY (1) 26+XY (2) 28+XY (1) 24+XY 26+XY Genus (number of species studied) Hahn, 1832 (1) Hahn, 28+XY Lepeletier and Serville, 1825