Isometopinae, Fieber, 1860
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publication ID |
https://doi.org/ 10.11646/zootaxa.5382.1.19 |
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publication LSID |
lsid:zoobank.org:pub:2D16527E-D679-4313-B46B-59B63CB813B5 |
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DOI |
https://doi.org/10.5281/zenodo.10288527 |
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persistent identifier |
https://treatment.plazi.org/id/047C87C9-FFDC-3E4D-FF01-F8CEFAE6FA21 |
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treatment provided by |
Plazi |
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scientific name |
Isometopinae |
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Monophyly of Isometopinae and position within Miridae
The Isometopinae Fieber, 1860 are a highly autapomorphic group possessing paired ocelli, which are absent in all other members of the plant bug family Miridae View in CoL ( Herczek 1993, Cassis & Schuh 2012, Namyatova & Cassis 2016, Yasunaga et al. 2017). This subfamily was considered to be the sister group to all other subfamilies based on morphology ( Schuh 1974, 1976), but recent works using molecular data do not support this hypothesis ( Schuh et al. 2009, Jung & Lee 2012).
Due to scarce information on habits, biology, and food preference, representatives are relatively rare in collections, with many species known from singletons or only a handful of specimens ( Eyles 1971, Namyatova & Cassis 2016, Taszakowski et al. 2020). For this reason, only a few representatives of this subfamily are included in phylogenetic studies if at all, and in the case of molecular datasets they are almost absent ( Schuh et al. 2009, Jung & Lee 2012, Kim & Jung 2019, Oh et al. 2023).
Morphology and character exploration
Isometopinae are a group of bugs with relatively small body size. Although the body length of Gigantometopus rossi Schwartz & Schuh, 1990 is 6.98 mm, most representatives are much smaller, and the body length ranges between 1.6 and 4 mm ( Schwartz & Schuh 1990, Akingbohungbe 1996). The most characteristic feature of Isometopinae is the possession of ocelli, which distinguishes them from other Miridae . Moreover, they are characterized by their often holoptic eyes, often strongly antero-posteriorly flattened head, simple phallotheca, a membranous endosoma with an undifferentiated secondary gonopore, pretarsal claws often with a subapical tooth and saltatorial hind legs ( Cassis & Schuh 2012, Henry 2017, Schuh & Weirauch 2020). An important feature of jumping tree bugs is the reduced number of femoral trichobothria, two and three on meso- and metafemora. The exceptions are Sophianini (three to four mesofemoral and four to five metafemoral trichobothria) and Gigantometopini (four to five mesofemoral and five to six metafemoral trichobothria) ( Schuh 1975, Akingbohungbe 1996, Yasunaga 2017, Taszakowski et al. 2021a, b).
The most comprehensive information on Isometopine morphology can be found in the studies by Herczek (1993) and Akingbohungbe (1996). A detailed description of male genitalia structure was provided by Konstantinov (2003). Female genitalia have not been the subject of comparative studies; we only know them from particular species descriptions ( Taszakowski et al. 2022).
Knowledge of the Isometopinae nymph’s morphology is very poor and limited to a few descriptions or photos included in papers. Recent work indicates that nymphs may have unusual characteristics comparing to those of the other subfamilies and require extensive study ( Yasunaga & Hayashi 2002, Yasunaga 2005, Yasunaga & Duwal 2006, Kim & Jung 2016, Çerçi & Dursun 2017, Yasunaga et al. 2017, Shishido et al. 2020, Kim et al. 2021, Yeshwanth et al. 2021).
Systematics and taxonomy of Isometopinae
Comprehensive information on the history of taxonomic and faunistic research on jumping tree bugs can be found in Herczek (1993). Since then, over 40 papers on jumping tree bugs have been published ( Fig. 1 View FIGURE 1 ). In the monographic study of the Isometopinae of Africa, Europe and the Middle East by Akingbohungbe (1996), the author described 37 new species. Following in the new century, in 2004 Lin described seven new species from Taiwan. Two years later, papers on jumping tree bugs from Yemen ( Akingbohungbe 2006) and Nepal ( Yasunaga & Duwal 2006) were published. Subsequently, in 2012, Henry & Carpintero reviewed the Isometopinae of Argentina and nearby areas of Brazil and Paraguay and described nine new species. In 2014, Herczek & Popov revised the genus Metoisops from late Eocene European amber, describing seven new species. Australian representatives of Isometopinae were the subject of research by Namyatova & Cassis (2016), who described seven new species and extensively discussed the distribution and host plant associations of the subfamily worldwide. In 2018, Krüger described ten new species from Liberia and three years later, Yeshwanth et al. (2021) revised the Isometopinae of India and Sri Lanka and described six new species. In summary, 152 species of Isometopinae have been described over the last 30 years, constituting over 50% of all known species.
Currently, six tribes, 53 genera and 289 species of Isometopinae are known, of which seven genera and 23 species are fossil taxa (see Checklist and Tab. 1 View TABLE 1 ).
The Diphlebini Bergroth, 1924 ( Fig. 2A View FIGURE 2 ) includes only a single genus, Diphleps Bergroth, 1924 with five known species. Although the systematic position of Diphlebini is debatable ( Herczek 1993, Akingbohungbe 1996, Konstantinov 2003, Cassis & Schuh 2012), we maintain this group in the Isometopinae , pending further study. Gigantometopini Herczek, 1993 ( Figs 2B View FIGURE 2 , 3A–C View FIGURE 3 ) is a small tribe containing nine genera and 27 species. The Isometopini Fieber, 1860 ( Figs 2C View FIGURE 2 , 3D–F View FIGURE 3 ) (14 genera, 127 species) and Myiommini Bergroth, 1924 ( Fig. 2D View FIGURE 2 ) (15 genera, 108 species) are the most species-rich isometopine tribes. Most recently, in 2017, Yasunaga et al. (2017) established the new tribe Sophianini Yasunaga, Yamada & Tsai, 2017 ( Figs 2E View FIGURE 2 , 3G, H View FIGURE 3 ) comprising two genera and ten species previously classified within Myiommini. The Electromyiommini Herczek, 1993 is an extinct tribe with four genera and eight species ( Fig. 4 View FIGURE 4 ).
The two genera, Isometopus Fieber, 1860 ( Fig. 2C View FIGURE 2 ) and Myiomma Puton, 1872 ( Fig. 2D View FIGURE 2 ), are distinguished by their high number of species, respectively 82 and 80. Representatives of these genera comprise 56% of all known jumping tree bugs. Isometopus and Myiomma probably are a ‘convenience group’ ( Namyatova & Cassis 2016). Redescription and diagnosis of these genera across its distribution range is undoubtedly a challenge waiting to be undertaken. Twenty-three genera of jumping three bugs are monotypic.
Checklist and zoogeography
The following checklist is based on the online catalog by Schuh (2002–2013). It also includes omitted species mentioned in the works of Linnavuori et al. (1998) and Akingbohungbe (2006) and the latest papers: Herczek & Popov (2011, 2012), Akingbohungbe (2012), Herczek et al. (2013, 2018, 2020), Kim & Jung (2016, 2021), Namyatova & Cassis (2016), Yasunaga et al. (2016, 2017), Çerçi & Dursun (2017), Hosseini (2017), Krüger (2018), Taszakowski et al. (2020, 2021a, b, 2022), Kim et al. (2021, 2023) and Yeshwanth et al. (2021).
Isometopus mahal Distant, 1911 was synonymized with I. mirificus Mulsant & Rey, 1879 by Carvalho (1951) but was treated as a valid species by Akingbohungbe (1996) and Yasunaga & Hayashi (2002). Fossil species were marked with a dagger (†). We have tried to review the available literature thoroughly, but there may be some omissions in the checklist. If you notice them, please kindly inform us.
The Wallace Line was adopted as the boundary between the Indomalayan and Australasian regions ( Rueda et al. 2013). The boundary of the Palearctic region was adopted according to Aukema & Rieger (1995). It is worth noting here that the Ryukyu Islands and Taiwan are placed in the Palearctic region; however, the fauna of these islands refers clearly to the Indomalayan fauna.
Explanation of color markings of zoogeographical regions: ●—Afrotropical, ●—Australasian, ●— Indomalayan, ●—Nearctic, ●—Neotropical, ●—Palearctic.
subfamily: Isometopinae Fieber, 1860
tribe: Diphlebini Bergroth, 1924
genus: Diphleps Bergroth, 1924
1. Diphleps henryi Hernandez, 1998 ●
2. Diphleps maldonadoi Henry, 1977 ●
3. Diphleps similaris Henry, 1977 ●
4. Diphleps unica Bergroth, 1924 ●
5. Diphleps yenli † Santiago-Blay & Poinar, 1993
tribe: Electromyiommini † Herczek, 1993
genus: Archemyiomma † Herczek, 1993
6. Archemyiomma carvalhoi † Herczek, 1993
7. Archemyiomma schaeferi † Herczek & Popov, 2013
genus: Clavimyiomma † Popov & Herczek, 1992
8. Clavimyiomma henryi † Popov & Herczek, 1992
genus: Electroisops † Herczek & Popov, 1997
9. Electroisops ritzkowskii † Herczek & Popov, 1997
genus: Electromyiomma † Popov & Herczek, 1992
10. Electromyiomma herczeki † Kim & Jung, 2021
11. Electromyiomma polonicum † Popov & Herczek, 1992
12. Electromyiomma schultzi † Popov & Herczek, 1992
13. Electromyiomma weitschati † Popov & Herczek, 1992
tribe: Gigantometopini Herczek, 1993
genus: Astroscopometopus Yasunaga & Hayashi, 2002
14. Astroscopometopus formosanus ( Lin, 2004) ●
15. Astroscopometopus gryllocephalus (Miyamoto, Yasunaga & Hayashi, 1996) ● 16. Astroscopometopus hesaraghattaensis Yeshwanth, Chérot & Henry, 2021 ● genus: Gigantometopus Schwartz & Schuh, 1990
17. Gigantometopus coronobtectus Kim, Taszakowski & Jung, 2021 ● 18. Gigantometopus rossi Schwartz & Schuh, 1990 ●
19. Gigantometopus schuhi Akingbohungbe, 2012 ●
genus: Isometopidea Poppius, 1913
20. Isometopidea lieweni Poppius, 1913 ●
21. Isometopidea viraktamathi Yeshwanth, Chérot & Henry, 2021 ●
genus: Kohnometopus Yasunaga, 2005
22. Kohnometopus fraxini Yasunaga, 2005 ●
23. Kohnometopus yangi (Lin, 2005) ●
24. Kohnometopus yasunagai Taszakowski, Kim & Masłowski, 2022 ●
genus: Megalofaciatus Taszakowski, Kim & Herczek, 2021
25. Megalofaciatus foliotibialis Taszakowski, Kim & Herczek, 2021 ●
26. Megalofaciatus gibbosus Taszakowski, Kim & Herczek, 2021 ● genus: Metoisops † Popov & Herczek, 1992
27. Metoisops akingbohungbei † Herczek & Popov, 2014
28. Metoisops consimilis † Herczek & Popov, 2014
29. Metoisops grabenhorsti † Herczek & Popov, 2014
30. Metoisops groehni † Herczek & Popov, 2014
31. Metoisops intergerivus † Herczek & Popov, 2014
32. Metoisops kerzhneri † Popov & Herczek, 1992
33. Metoisops michalskii † Kim, Taszakowski & Herczek, 2023
34. Metoisops popovi † Kim, Taszakowski & Jung, 2023
35. Metoisops punctatodiffusus † Herczek & Popov, 2014
36. Metoisops punctatus † Popov & Herczek, 1993
37. Metoisops variabilis † Herczek & Popov, 2014
genus: Sulawesimetopus Herczek, Gorczyca & Taszakowski, 2018
38. Sulawesimetopus henryi Herczek, Gorczyca & Taszakowski, 2018 ● genus: Planicapitus Taszakowski, Kim & Herczek, 2020
39. Planicapitus luteus Taszakowski, Kim & Herczek, 2020 ●
genus: Bruneimetopus Taszakowski, Kim & Herczek, 2020
40. Bruneimetopus simulans Taszakowski, Kim & Herczek, 2020 ● tribe: Isometopini Fieber, 1860
subtribe: Isometopina Fieber, 1860
genus: Carayonischa Akingbohungbe, 1996
41. Carayonischa singularis Akingbohungbe, 1996 ●
genus: Eurocrypha Kirkaldy, 1908
42. Eurocrypha thanatochlamys Kirkaldy, 1908 ●
genus: Isometopiellus Akingbohungbe, 1996
43. Isometopiellus cylpoides Akingbohungbe, 1996 ●
44. Isometopiellus heterocephalus (Puton, 1898) ●
45. Isometopiellus palliceps (Wagner, 1973) ●
46. Isometopiellus ugandanus Akingbohungbe, 1996 ●
genus: Isometopus Fieber, 1860
47. Isometopus africanus Herczek, 2004 ●
48. Isometopus albifrons (Slater & Schuh, 1969) ●
49. Isometopus amurensis Kerzhner, 1988 ●
50. Isometopus angolensis Hoberlandt, 1952 ●
51. Isometopus anlasi Çerçi & Dursun, 2017 ●
52. Isometopus aureus Akingbohungbe, 1996 ●
53. Isometopus beijingensis Ren & Yang, 1988 ●
54. Isometopus bongensis Krüger, 2018 ●
55. Isometopus bipunctatus Lin, 2004 ●
56. Isometopus brevirostris Akingbohungbe, 1996 ●
57. Isometopus chaiyaphum Yasunaga, Duanthisan & Yamada, 2016 ●
58. Isometopus carnifrons Akingbohungbe, 2006 ●
59. Isometopus citri Ren, 1987 ●
60. Isometopus confusus (Akingbohungbe, 1983) ●
61. Isometopus cuneatus (Distant, 1904) ●
62. Isometopus deemingi (Akingbohungbe, 1983) ● 63. Isometopus discrepans Akingbohungbe, 1996 ● 64. Isometopus diversiceps Linnavuori, 1962 ●
65. Isometopus fallax Akingbohungbe, 1996 ●
66. Isometopus fasciatus Hsiao, 1964 ●
67. Isometopus frontalis Akingbohungbe, 1996 ● 68. Isometopus fulvus (Akingbohungbe, 1983) ● 69. Isometopus gharaati Akingbohungbe, 2012 ● 70. Isometopus hainanus Hsiao, 1964 ●
71. Isometopus hananoi Hasegawa, 1946 ●
72. Isometopus hasegawai Miyamoto, 1965 ●
73. Isometopus insperatus Akingbohungbe, 1996 ● 74. Isometopus intermedius Akingbohungbe, 1996 ● 75. Isometopus intrusus (Herrich-Schaeffer, 1835) ● 76. Isometopus ishigaki Yasunaga, 2005 ●
77. Isometopus japonicus Hasegawa, 1946 ●
78. Isometopus jejuensis Kim & Jung, 2016 ●
79. Isometopus kanako Yasunaga & Duwal, 2006 ● 80. Isometopus kaznakovi Kiritshenko, 1939 ●
81. Isometopus lineatifrons Akingbohungbe, 1996 ● 82. Isometopus lini Lin, 2004 ●
83. Isometopus linnavuorii Hosseini, 2017 ●
84. Isometopus longirostris Akingbohungbe, 1996 ● 85. Isometopus longisetosus ( Herczek, 1993) ●
86. Isometopus lunaris Linnavuori, 1975 ●
87. Isometopus maculipennis Akingbohungbe, 2004 ● 88. Isometopus maculosus Akingbohungbe, 1996 ● 89. Isometopus madagascariensis (Polhemus, 1988) ● 90. Isometopus mahal (Distant, 1911) ●
91. Isometopus marginatus Ren & Yang, 1988 ● 92. Isometopus mirificus Mulsant & Rey, 1879 ● 93. Isometopus mirus Akingbohungbe, 2004 ●
94. Isometopus nagarjun Yasunaga & Duwal, 2006 ● 95. Isometopus nigrans Akingbohungbe, 1996 ● 96. Isometopus nigritulus Akingbohungbe, 1996 ● 97. Isometopus nigrosignatus Ren, 1987 ●
98. Isometopus nitidus (Smith, 1967) ●
99. Isometopus obesulus Akingbohungbe, 1996 ● 100. Isometopus peltatus McAtee & Malloch, 1932 ● 101. Isometopus peregrinus Akingbohungbe, 1996 ● 102. Isometopus pictus Linnavuori, 1975 ●
103. Isometopus praetermissum Akingbohungbe, 2012 ● 104. Isometopus puberus Ren, 1991 ●
105. Isometopus puncticollis Akingbohungbe, 1996 ● 106. Isometopus quadrifasciatus Wagner, 1973 ● 107. Isometopus quadrivittatus Akingbohungbe, 2003 ● 108. Isometopus renae Lin, 2004 ●
109. Isometopus rugiceps Kerzhner, 1988 ●
110. Isometopus sepehrii Linnavuori, Sarafrazi & Hosyni, 1998 ● 111. Isometopus shaowuensis Ren, 1987 ●
112. Isometopus siamensis Yasunaga & Yamada 2013 ● 113. Isometopus slateri (Akingbohungbe, 1983) ●
114. Isometopus sudanicus Akingbohungbe, 2006 ●
115. Isometopus taeniaticeps Puton, 1898 ●
116. Isometopus takaii Yasunaga, 2005 ●
117. Isometopus tianjinus Hsiao, 1964 ●
118. Isometopus tibialis (Akingbohungbe, 1983) ●
119. Isometopus transvaalensis (Slater & Schuh, 1969) ● 120. Isometopus turneri (Slater & Schuh, 1969) ●
121. Isometopus typicus (Distant, 1910) ●
122. Isometopus vanharteni Akingbohungbe, 2006 ●
123. Isometopus variabilis Akingbohungbe, 1996 ●
124. Isometopus wacriensis (Smith, 1967) ●
125. Isometopus webbi Yeshwanth, Chérot & Henry, 2021 ● 126. Isometopus wolskii Yeshwanth, Chérot & Henry, 2021 ● 127. Isometopus yemenensis Akingbohungbe, 2003 ● 128. Isometopus yoshizawai Yasunaga & Duwal, 2006 ●
genus: Lindbergiola Carvalho, 1951
129. Lindbergiola aureopilosa Carvalho, 1951 ●
130. Lindbergiola bicolor (McAtee & Malloch, 1932) ● 131. Lindbergiola jarmilae Hoberlandt, 1952 ●
genus: Paloniella Poppius, 1915
132. Paloniella annulata (Ren & Huang, 1987) ●
133. Paloniella bedfordi (Hesse, 1947) ●
134. Paloniella cuneata (Slater & Schuh, 1969) ●
135. Paloniella erinacea Krüger, 2018 ●
136. Paloniella feana (Distant, 1904) ●
137. Paloniella flavicolor Akingbohungbe, 2004 ●
138. Paloniella garmsi Krüger, 2018 ●
139. Paloniella latifrons Akingbohungbe, 1996 ●
140. Paloniella microchelys Yasunaga, Duanthisan & Yamada, 2016 ● 141. Paloniella montana (Ren & Yang, 1988) ●
142. Paloniella mutabilis Akingbohungbe, 1996 ●
143. Paloniella niger (Linnavuori, 1975) ●
144. Paloniella nodifrons Akingbohungbe, 2003 ●
145. Paloniella ovata Akingbohungbe, 2006 ●
146. Paloniella parallela Yasunaga & Hayashi, 2002 ●● 147. Paloniella pellucida Akingbohungbe, 1996 ●
148. Paloniella pseudotyloides Akingbohungbe, 1996 ● 149. Paloniella senegalensis Akingbohungbe, 1996 ●
150. Paloniella suffuscipennis Akingbohungbe, 1996 ● 151. Paloniella tafoensis Akingbohungbe, 1996 ●
152. Paloniella umbrosa (Slater & Schuh, 1969) ●
153. Paloniella xizangana (Ren, 1988) ●
genus: Ptisca McAtee & Malloch, 1932
154. Ptisca blattiformis McAtee & Malloch, 1932 ●
155. Ptisca liberiense Krüger, 2018 ●
genus: Smithopus Akingbohungbe, 1996
156. Smithopus ghanaiensis (Smith, 1967) ●
157. Smithopus scutellaris (Linnavuori, 1975) ●
subtribe: Nesocryphina Herczek, 1993
genus: Australotopus Namyatova & Cassis, 2016
158. Australotopus cooperensis Namyatova & Cassis, 2016 ●
genus: Fronsonia Herczek, 1993
159. Fronsonia ochracea Herczek, 1993 ●
genus: Jozefus Herczek, 1993
160. Jozefus brunetus Namyatova & Cassis, 2016 ●
161. Jozefus guineiensis Herczek, 1993 ●
162. Jozefus monteithi Namyatova & Cassis, 2016 ●
genus: Nesocrypha Kirkaldy, 1908
163. Nesocrypha corticicola Kirkaldy, 1908 ●
genus: Paratopus Herczek, 1993
164. Paratopus brunocapitus Namyatova & Cassis, 2016 ●
165. Paratopus flavocapitus Namyatova & Cassis, 2016 ●
166. Paratopus ovatus (Herczek, 1991) ●
genus: Popoviana Herczek & Popov, 1997
167. Popoviana fijiensis ( Herczek, 1993) ●
tribe: Myiommini Bergroth, 1924
genus: Namaquaropus Akingbohungbe, 2004
168. Namaquaropus niger Akingbohungbe, 2004 ●
subtribe: Myiommina Bergroth, 1924
genus: Bongiella Krüger, 2018
169. Bongiella nodistylis Krüger, 2018 ●
genus: Brailovskiocoris Henry, 1980
170. Brailovskiocoris nocturnus (Brailovsky, 1976) ●●
genus: Corticoris McAtee & Malloch, 1922
171. Corticoris infuscatus Henry & Herring, 1979 ●●
172. Corticoris libertus (Gibson, 1917) ●
173. Corticoris mexicanus Henry & Herring, 1979 ●
174. Corticoris pallidus Henry, 1984 ●
175. Corticoris pintoi Henry, 1984 ●
176. Corticoris pubescens Henry, 1984 ●
177. Corticoris pulchellus (Heidemann, 1908) ●
178. Corticoris signatus (Heidemann, 1908) ●
179. Corticoris unicolor (Heidemann, 1908) ●
genus: Lidopiella Henry, 1980
180. Lidopiella slateri Henry, 1980 ●
genus: Lidopus Gibson, 1917
181. Lidopus heidemanni Gibson, 1917 ●
182. Lidopus schwarzi ( McAtee & Malloch, 1924) ●
genus: Myiomma Puton, 1872
183. Myiomma adusta Herczek, 2004 ●
184. Myiomma affinis (Hoberlandt, 1952) ●
185. Myiomma albalata Namyatova & Cassis, 2016 ●
186. Myiomma albicoxa Smith, 1967 ●
187. Myiomma albiscutellata Smith, 1967 ●
188. Myiomma albostiolata Krüger, 2018 ●
189. Myiomma altica Ren, 1987 ●
190. Myiomma amaranion Herczek & Popov, 2006 ●
191. Myiomma apicalis Henry & Carpintero, 2012 ●
192. Myiomma argentinensis Henry & Carpintero, 2012 ●
193. Myiomma austroccidens Yasunaga, Yamada & Tsai, 2017 ●
194. Myiomma basseti Namyatova & Cassis, 2016 ●
195. Myiomma belavadii Yeshwanth, Chérot & Henry, 2021 ●
196. Myiomma bionotata Henry & Carpintero, 2012 ●
197. Myiomma brasilianum Henry, 1979 ●
198. Myiomma bredoi Akingbohungbe, 1996 ●
199. Myiomma brunnea Krüger, 2018 ●
200. Myiomma capeneri Slater & Schuh, 1969 ●
201. Myiomma capitatum Henry, 1979 ●
202. Myiomma choui Lin & Yang, 2004 ●
203. Myiomma cixiiforme (Uhler, 1891) ●
204. Myiomma cobbeni Akingbohungbe, 2003 ●
205. Myiomma confusa Akingbohungbe, 1996 ●
206. Myiomma dundoensis (Hoberlandt, 1952) ●
207. Myiomma fasciata Smith, 1967 ●
208. Myiomma ferruginea Akingbohungbe, 1996 ●
209. Myiomma fieberi Puton, 1872 ●
210. Myiomma fulva Smith, 1967 ●
211. Myiomma fuscipes Krüger, 2018 ●
212. Myiomma fusiforme Henry, 1979 ●
213. Myiomma goellneri Krüger, 2018 ●
214. Myiomma hemialba ( Carvalho, 1951) ●
215. Myiomma impunctata Smith, 1967 ●
216. Myiomma jankotejai Herczek & Popov, 2006 ● 217. Myiomma juniperina Linnavuori, 1975 ●
218. Myiomma keltoni Henry, 1984 ●
219. Myiomma kentingense Yasunaga, Yamada & Tsai, 2017 ● 220. Myiomma kukai Yasunaga & Hayashi, 2002 ●
221. Myiomma lansburyi ( Carvalho, 1951) ●
222. Myiomma latifrons Herczek, 2004 ●
223. Myiomma linearis Akingbohungbe, 1996 ●
224. Myiomma lutea McAtee & Malloch, 1932 ●
225. Myiomma maculata Akingbohungbe, 2003 ●
226. Myiomma mexicanum Henry, 1979 ●
227. Myiomma milleri (Hoberlandt, 1959) ●
228. Myiomma minor Akingbohungbe, 1996 ●
229. Myiomma minutum Miyamoto, 1965 ●
230. Myiomma montana Linnavuori, 1975 ●
231. Myiomma nigricole Akingbohungbe, 2006 ●
232. Myiomma nigra Smith, 1967 ●
233. Myiomma obscura Akingbohungbe, 1996 ●
234. Myiomma ornatum Henry, 1979 ●
235. Myiomma ostentans Akingbohungbe, 1996 ●
236. Myiomma pallidopleura Henry & Carpintero, 2012 ● 237. Myiomma pallipes Henry & Carpintero, 2012 ● 238. Myiomma phuvasae Yasunaga, Duanthisan & Yamada, 2016 ● 239. Myiomma piceicola Akingbohungbe, 1996 ●
240. Myiomma qinlingensis Qi, 2005 ●
241. Myiomma ramamurthyi Yeshwanth, Chérot & Henry, 2021 ● 242. Myiomma rubida Akingbohungbe, 1996 ●
243. Myiomma rubra Smith, 1967 ●
244. Myiomma rubrooculatum Henry, 1979 ●
245. Myiomma rubrovenata Smith, 1967 ●
246. Myiomma samuelsoni Miyamoto, 1965 ●
247. Myiomma schmitzi Slater, 1976 ●
248. Myiomma schuhi Henry, 1979 ●
249. Myiomma scotti Herczek, 2004 ●
250. Myiomma scutellata Henry & Carpintero, 2012 ● 251. Myiomma semipallidum Henry, 1979 ●
252. Myiomma surinamensis (Carvalho & Rosas, 1962) ● 253. Myiomma takahashii Yasunaga & Hayashi, 2002 ● 254. Myiomma uniformis Henry & Carpintero, 2012 ● 255. Myiomma ussuriensis Ostapenko, 2001 ●
256. Myiomma variabilis Krüger, 2018 ●
257. Myiomma verticata Smith, 1967 ●
258. Myiomma vittata McAtee & Malloch, 1932 ●
259. Myiomma vittaticornis Akingbohungbe, 1996 ●
260. Myiomma voigti † ( Popov & Herczek, 1992)
261. Myiomma zandeana Linnavuori, 1975 ●
262. Myiomma zhengi Lin & Yang, 2004 ●
genus: Myiopus Henry, 1980
263. Myiopus woldai Henry, 1980 ●
genus: Slateropus Akingbohungbe, 1996
264. Slateropus miriformis (Slater & Schuh, 1969) ● 265. Slateropus perplexus Akingbohungbe, 1996 ●
genus: Wetmorea McAtee & Malloch, 1924
266. Wetmorea notabilis McAtee & Malloch, 1924 ●
subtribe: Plaumannocorina Herczek, 1993
genus: Aristotelesia Carvalho, 1947
267. Aristotelesia carioca Carvalho, 1947 ●
268. Aristotelesia fuscata Henry & Carpintero, 2012 ● 269. Aristotelesia medialis Henry & Carpintero, 2012 ●
genus: Joceliana Carvalho, 1984
270. Joceliana graziae Carvalho, 1984 ●
genus: Plaumannocoris Carvalho, 1947
271. Plaumannocoris rarus Carvalho, 1947 ●
subtribe: Tottina Herczek, 1993
genus: Paratotta Herczek, 1993
272. Paratotta orientalis Herczek, 1993 ●●
genus: Totta Ghauri & Ghauri, 1983
273. Totta puspae Yasunaga & Duwal, 2006 ●
274. Totta rufercorna Lin & Yang, 2004 ●
275. Totta zaherii Ghauri & Ghauri, 1983 ●
tribe: Sophianini Yasunaga, Yamada & Tsai, 2017
genus: Alcecoris McAtee & Malloch, 1924
276. Alcecoris fraxinusae Lin, 2004 ●
277. Alcecoris formosanus Lin, 2004 ●
278. Alcecoris globosus Carvalho, 1951 ●
279. Alcecoris lamellatus (Ren & Yang, 1988) ●
280. Alcecoris linyangorum Yasunaga, Yamada & Tsai, 2017 ● 281. Alcecoris cochlearatus Yasunaga, Yamada & Tsai, 2017 ● 282. Alcecoris periscopus McAtee & Malloch, 1924 ●● 283. Alcecoris heissi Herczek & Popov, 2011 ●
genus: Sophianus Distant, 1904
284. Sophianus alces Distant, 1904 ●
285. Sophianus kerzhneri Lin, 2009 ●
286. Sophianus palawanensis Taszakowski, Kim & Herczek, 2021 ● Incertae sedis genus: Hoffheinsoria † Herczek & Popov, 2012
287. Hoffheinsoria robusta † Herczek & Popov, 2012
genus: Isomyiomma † Herczek, Popov & Drohojowska, 2020
288. Isomyiomma hirta † Herczek, Popov & Drohojowska, 2020
genus: Sagarmathametopus Yasunaga & Duwal, 2006
289. Sagarmathametopus fuscescens Yasunaga & Duwal, 2006 ●
Isometopinae is a group inhabiting mainly the Old World, from which 222 out of 266 extant species are known (83%). The greatest species diversity occurs in the Afrotropical region—95 species. Seventy-four species are known from the Palearctic region, 40 from the Indomalayan region and 19 from the Australasian region. The New World is represented by only 44 species, 30 in the Neotropical region and 14 in the Nearctic region.
Over the last 30 years, significant progress has been achieved in studying the fauna of the Afrotropical, Palearctic, Australasian, and especially the Indomalayan region (67% of known species from the latter area have been described). Relatively little attention has been paid to the New World Isometopinae . Ten Neotropical species (Hernandez 1998, Henry & Carpintero 2012) and no Nearctic species were described.
The occurrence of higher taxa in particular zoogeographic regions is presented in Table 1 View TABLE 1 . Diphlebini are a group limited in range to the New World. Gigantometopini and Sophianini occur in Australasian and Indomalayan regions, as well as warm areas of the Palearctic. Representatives of Isometopini are known from the Old World, while Myiommini are the most widely distributed tribe of jumping tree bugs and are characterized by a cosmopolitan range. There is no modern analysis of the distribution of Isometopinae . Discussion of distribution was presented by Herczek (1993) and Namyatova & Cassis (2016).
Fossils
Isometopinae live on the bark of trees ( Namyatova & Cassis 2016, Yasunaga 2017), which predisposes them, like Cylapinae ( Wolski 2021) , to be more easily fossilized in resins than the other groups of Miridae , given that most of the described fossils are cylapines and isometopines ( Schuh & Weirauch, 2020). Nevertheless, the first species of fossil representative of jumping tree bugs was described just over 30 years ago ( Popov & Herczek 1992). As we already mentioned, seven genera and 23 species of Isometopinae are of fossil taxa ( Popov & Herczek 2008; Herczek & Popov 2012, 2014; Herczek et al. 2013, 2020; Kim & Jung 2021; Kim et al. 2023). Eight species classified into four genera belong to the entirely fossil tribe Electromyiommini ( Fig. 4A, B, D, E View FIGURE 4 ). Until recently, the aforementioned tribe also included the genus Metoisops , which, with 11 species, is the most numerous of the fossil genera of Isometopinae . In 2023, Kim et al. transferred the Metoisops to the Gigantometopini. Two extant genera, Diphleps and Myiomma , have one fossil representative each. In addition, it is herein noticed that two species, Hoffheinsoria robusta ( Fig. 4C View FIGURE 4 ) and Isomyiomma hirta , currently have incertae sedis status. Except Diphleps yenli , which is known from Miocene Dominican amber ( Santiago-Blay & Poinar 1993), all other fossil representatives of jumping tree bugs come from Eocene Baltic amber ( Popov & Herczek 2008; Herczek & Popov 2012, 2014; Herczek et al. 2013, 2020; Kim & Jung 2021; Kim et al. 2023).
The incertae sedis taxa and the genera included in Electromyiommini certainly require further research regarding their systematic position ( Kim et al. 2023).
Biology
Jumping tree bugs have a cryptic habit, which results in rare observations in their natural environment. They are typically collected on tree bark or in low light and damp conditions ( Akingbohungbe 1996, Yeshwanth et al. 2021). The few papers that mention the biology of Isometopinae indicate that they are zoophagous, feeding commonly on scales ( Wheeler & Henry 1978, Ghauri & Ghauri 1983, Henry 1984, Akingbohungbe 1996, Wheeler 2001). However, this information only applies to a few species in the genera Diphleps (Diphlebini) , Isometopus and Paloniella (Isometopini) and Corticoris , Myiomma & Lidopus (Myiommini) . We still have no knowledge of the trophic relationships of representatives of Gigantometopini and Sophianini . Photos of representatives of these tribes (and other isometopines), included in various papers (e.g., Kim & Jung 2016; Yeshwanth et al. 2021) as well as images increasingly available on the Internet, show bugs that appear to feed on lichen growing on tree bark ( Fig. 3 View FIGURE 3 ). These observations confirm previous suggestions of several authors ( McAtee & Malloch 1924, Schuh 1976). The high quality of the abovementioned photos enables us to see what the bugs feed; however, there are no visible arthropods. It may be conceivable that feeding on scales is opportunistic feeding due to the fact that scales are immobile, and phytophagous insects (including plant bugs) sometimes feed on the other arthropods ( Cobben 1978, Eubanks et al. 2003). In addition, previous phylogenetic studies hypothesized the close relationship between Isometopinae and Cylapinae ( Schuh et al. 2009, Oh et al. 2023). Given that relatively small groups within Miridae (e.g., Bryocorinae , Cylapinae , and Isometopinae ) have more specific feeding habits, and the cylapines are well known as to a large extent fungivorous group ( Gossner & Damken 2018, Kim et al. 2019, Wolski 2021), it is highly possible that the isometopines are lichen feeder and not strictly zoophagous.
Namyatova & Cassis (2016) presented a discussion of the distribution and host plant associations of the subfamily on a worldwide basis. Trees or large shrubs are most plant records for Isometopinae . Nevertheless, few species are known from small shrubs, vines or herbs. The greatest number of species was recorded from angiosperms.
TABLE 1. Summary of abundance and zoogeography of higher taxa of extant Isometopinae.Explanation of color markings of zoogeographical regions: ●—Afrotropical, ●—Australasian, ●—Indomalayan, ●—Nearctic, ●—Neotropical, ●— Palearctic.
| tribe | number of genera | number of species | subtribe | number of genera | number of species |
|---|---|---|---|---|---|
| Diphlebini ●● | 1 | 4 | - | ||
| Gigantometopini ●●● | 8 | 16 | - | ||
| Isometopini ●●●● | 14 | 127 | Isometopina ●●●● Nesocryphina ● | 8 6 | 117 10 |
| Myiommina ●●●●●● | 9 | 97 | |||
| Myiommini ●●●●●● | 15 | 107 | Plaumannocorina ● Tottina ●●● | 3 2 | 5 4 |
| incertae sedis ● | 1 | 1 | |||
| Sophianini ●●● | 2 | 11 | - | ||
| incertae sedis ● | 1 | 1 | - | ||
| Together | 41 | 266 |
TABLE 1. Summary of abundance and zoogeography of higher taxa of extant Isometopinae . Explanation of color markings of zoogeographical regions: ●—Afrotropical, ●—Australasian, ●—Indomalayan, ●—Nearctic, ●—Neotropical, ●— Palearctic.
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