On the morphology and taxonomy of the genera Halys and Neohalys (Heteroptera: Pentatomidae: Halyini)

D.A. Gapon

Abstract. The male and female terminalia of Halys sulcatus (Thunberg, 1783) and H. magnus Chopra, 1974, nom. resurr. are redescribed in detail, including the completely inflated aedeagi. The distribution of these taxa is analysed. It is shown that H. sulcatus is reliably recorded only from the southern regions of India and H. magnus, only from the northern regions; numerous published records of “H. dentatus” from the central regions of India cannot be reliably attributed to either of these taxa. It is shown that almost all characters of the terminalia previously used to distinguish between H. sulcatus and H. magnus are highly variable and do not have a hiatus in their variability. Furthermore, the specimens were examined which exhibited an intermediate type of paramere between those in the two taxa. The material examined (which is not very extensive) revealed only one character that can consistently differentiate between northern and southern populations: the shape of armament on the anterior margin of parameral hypophysis (i.e., the position of a denticle and the presence or absence of a carina proximal to its base). However, a lack of correlation between these characters and with other, even less significant characters, as well as published evidence of the variability of the paramere, raise doubts about whether H. sulcatus and H. magnus are distinct species. Hypotheses about the status of these taxa are considered. It is shown that the name H. sulcatus was applied to the taxon described as H. magnus, based on a variable character of the female terminalia, which has no diagnostic value in this case. Additionally, the type locality of H. sulcatus is in the southern regions of India, where no reliably identified specimens of H. magnus have been found so far. Therefore, in this article H. sulcatus is recognised as a senior synonym of H. serriger Westwood, 1837, syn. resurr., and the name H. magnus Chopra, 1974, nom. resurr., is resurrected to the taxon distributed in the north. In addition, a detailed redescription of the female terminalia of Halys shaista Ghauri, 1988 and the male and female terminalia of Neohalys serricollis (Westwood, 1837) is provided. A new species, Halys brocchus sp. nov. is described, distinguished from all congeners primarily by a unique character of its pygophore. An updated identification key for species of the genus Halys Fabricius, 1803 is given. The name H. hyderabadiensis Memon, Parveen, Ahmad et Shah, 2017, syn. nov. is recognised as a junior synonym of H. magnus. Neohalys Azim, 2002, syn. nov., is recognised as a junior subjective synonym and a junior homonym of Neohalys Ahmad et Perveen, 1982. The names Neohalys acuticornis Ahmad et Perveen, 1982, syn. nov., N. longirostratus Ahmad et Perveen, 1982, syn. nov., and N. minirostratus Ahmad et Perveen, 1982, syn. nov. are placed in synonymy with N. serricollis.

Key words: taxonomy, redescription, variability, male and female terminalia, completely inflated aedeagus, India, South Asia, Heteroptera, Pentatomidae, Halyini, new synonyms, new species

Zoosystematica Rossica, 2023, 32(2): 342–383 ▪ Published online 31 December 2023

https://doi.org/10.31610/zsr/2023.32.2.342 ▪ Open full article

References

Abbasi Q.A. 1986. Morpho-taxonomic studies of the family Pentatomidae Leach, 1815 (Heteroptera: Pentatomorpha) of South Asia (Pakistan, Azad Kashmir and Bangladesh) with reference to the phylogeny of the group. Pakistan Journal of Entomology, Supplement 5: 1–279.

Abbasi Q.A. & Ahmad I. 1976. A new species of the genus Halys Fabr. (Pentatomidea: Halyini) from East Bengal with notes on its genitalia and their bearing on classification. Karachi University Journal of Science, 4(1–2): 26–31.

Ahmad I. 1977. Systematics and biology of pentatomomorphous superfamilies Coreoidea and Pentatomoidea of Pakistan. Research Project No. FG-Pa-181 (A-17-ENT-37). Final report. Karachi: University of Karachi. vii + 625 p.

Ahmad I., Abbasi Q.A. & Khan A.A. 1974. Generic and supergeneric keys with reference to a check list of pentatomid fauna of Pakistan (Heteroptera: Pentatomoidea) with notes on their distribution and food plants. Proceedings of the Entomological Society of Karachi, Supplement 1: 1–103.

Ahmad I., Memon N. & Parveen R. 1998. A revision of the genus Neohalys Ahmad & Parveen (Heteroptera: Pentatomidae: Pentatominae) based on type material. Pakistan Journal of Entomology, 13(1–2): 1–4.

Ahmad I. & Parveen R. 1982. A new genus and three new species of Halyini from Pakistan with a note on their relationships (Hemiptera: Heteroptera: Pentatomidae). Transactions of the Shikoku Entomological Society, 16(1–2):1–10.

Ahmed N. & Bajwa G.A. 2016. Hemiptera fauna of Haripur District, Pakistan. Pakistan Entomologist, 38(1): 43–45.

Atkinson E.T. 1884. The Himalayan districts of the north-western provinces of India, 2. Allahabad: North-Western Provinces and Oudh Government Press. xviii + 964 p.

Azim M.N. 2002. Studies on Indian genera of the tribe Halyini (Pentatomidae: Pentatominae). Oriental Science, 2002: 41–52.

Azim M.N. 2011. Taxonomic survey of stink bugs (Heteroptera: Pentatomidae) of India. Halteres, 3: 1–10.

Biswas B. & Bal A. 2007. Insecta: Hemiptera: Heteroptera: Pentatomoidea. In: State fauna series, 5. Fauna of Andhra Pradesh (Part 3). Insects: 301–320. Kolkata: Zoological Survey of India.

Biswas B. & Bal A. 2010. Insecta: Hemiptera: Pentatomoidea. In: State fauna series, 18. Fauna of Uttarakhand (Part 2). Insects: 229–244. Kolkata: Zoological Survey of India.

Biswas B. & Ghosh L.K. 1995. Insecta: Hemiptera. In: Fauna of conservation areas, 6. Fauna of Indravati Tiger Reserve, Madhya Pradesh: 19–29. Calcutta: Zoological Survey of India.

Biswas B., Hassan M.E., Chandra K., Kushwaha S. and Mukherjee P. 2014. On an account of Pentatomoidea (Heteroptera: Hemiptera) from Chhattisgarh, India. Records of the Zoological Survey of India, 114(2): 211–231.

Chakraborty S.P. & Ghosh L.K. 1998. Insecta: Hemiptera: Heteroptera: Pentatomoidea. In: State fauna series, 4. Fauna of Meghalaya, 4: 391–429. Calcutta: Zoological Survey of India.

Chakraborty S.P. & Ghosh L.K. 2003. Insecta: Hemiptera: Heteroptera: Pentatomoide [sic!]. In: State fauna series, 9. Fauna of Sikkim, 2: 509–524. Kolkata: Zoological Survey of India.

Chakraborty S.P., Ghosh L.K. & Basu R.C. 1994. Insecta: Hemiptera: Pentatomoidea – 1. In: State fauna series, 3. Fauna of West Bengal, 5: 469–493. Calcutta: Zoological Survey of India.

Chandra A. 1953. Part 14. – Order Hemiptera (in part). Family Pentatomidae. In: Indian forest Leaflet, 121(3). Systematic catalogue of the main identified entomological collection at the Forest Research Institute, Dehra Dun, Part 14: 83–102.

Chandra K. 2008. Insecta Hemiptera. In: Faunal diversity of Jabalpur District, Madhya Pradesh: 141–157. Kolkata: Zoological Survey of India.

Chandra K. 2009. Insecta: Hemiptera. In: Conservation area series, 39. Fauna of Pachmarhi Biosphere Reserve: 247–257. Kolkata: Zoological Survey of India.

Chandra K., Biswas B. & Kushwaha S. 2014. Superfamily Pentatomoidea (Hemiptera: Heteroptera) of Madhya Pradesh, India. Munis Entomology & Zoology, 9(1): 198–205.

Chandra K., Bhandari R. & Kushwaha S. 2015a. Hemiptera fauna of Ratapani wildlife sanctuary, Madhya Pradesh, India. International Journal of Fauna and biological Sciences, 2(2): 20–26.

Chandra K., Bhandari R. & Kushwaha S. 2015b. On a collection of true bugs (Insecta: Hemiptera) from Ralamandal Wildlife Sanctuary, Madhya Pradesh, India. Munis Entomology & Zoology, 10(2): 486–488.

Chandra K. & Kushwaha S. 2013. Distribution and diversity of Hemiptera fauna of Singhori Wildlife Sanctuary, Raisen District, Madhya Pradesh, India. Munis Entomology & Zoology, 8(2): 677–681.

Chandra K., Kushwaha S., Sambath S. & Biswas B. 2012. Distribution and diversity of Hemiptera fauna of Veerangana Durgavati Wildlife Sanctuary, Damoh, Madhya Pradesh (India). Biological Forum –
an international Journal, 4(1): 68–74.

Chandra K. & Rajan P.T. 2004. Faunal Diversity of Mount Harriet National Park (South Andaman). Zoological Survey of India, Conservation Area Series, 17: 1–142. Kolkata: Zoological Survey of India.

Chopra N.P. 1972. A new species of the genus Agaeus Dallas (Hemiptera: Pentatomidae). Oriental Insects, 6(3): 305–307. https://doi.org/10.1080/00305316.1972.10434080

Chopra N.P. 1974. Studies on the genus Halys (Hemiptera: Heteroptera: Pentatomidae). Oriental Insects, 8(4): 473–479. https://doi.org/10.1080/00305316.1974.10434879

Dallas W.S. 1851. List of the specimens of hemipterous insects in the collection of the British Museum, 1. London: Trustees of the British Museum. 368 p. + 11 pls.

Dhiman S.C. & Yadav Y.K. 2005. Some ethological aspects of Halys dentatus Fabr. (Heteroptera: Pentatomidae). Journal of applied zoological Research, 16(1): 44–47.

Dhiman S.C. & Yadav Y.K. 2006. Oviposition behaviour and egg structure of Halys dentatus Fabr. a pest of Azadirachta indica A. Juss. Advances in Indian Entomology: Productivity & Health, 2: 23–32.

Dhiman S.C. & Yadav Y.K. 2008. Life history of a phytosuccivorus bug, Halys dentatus Fabr. (Heteroptera: Pentatomidae), a pest of neem tree, Azadirachta indica A. Juss, in western Uttar Pradesh. Indian Forester, 134(7): 958–976.

Distant W.L. 1902. The Fauna of British India, including Ceylon and Burma. Published under the authority of the secretary of state for India in council, 1. Heteroptera. London: W.T. Blanford. xxxviii + 438 p.

Dupuis C. 1955. Les genitalia des Hémiptères Hétéroptères: genitalia externes des deux sexes. Voies ectodermiques femelles. Mémoires du Muséum national d’Histoire naturelle,Série A, Zoologie, 6(4): 183–278.

Gaikwad S.M. & Waghmare S.H. 2018. Stink bugs (Hemiptera: Heteroptera: Pentatomidae) and plant association in Kolhapur district of Northern Western Ghats. Journal of emerging Technologies and innovative Research, 5(12):12–15.

Gapon D.A. 2001. Inflation of Heteropteran aedeagi using microcapillaries (Heteroptera, Pentatomidae). Zoosystematica Rossica, (2000), 9(1): 157–160.

Gapon D.A. 2008a. A revision of Leprosoma Baerensprung, 1859 (Heteroptera: Pentatomidae). In: Michail Josifov Festschrift. Advances in Heteroptera research. Festschrift in honor of 80th anniversary of Michail Josifov: 105–120. Sofia – Moscow: Pensoft Publishers.

Gapon D.A. 2008b. New subtribes and a new genus of Podopini (Heteroptera: Pentatomidae: Podopinae). Acta entomologica Musei Nationalis Pragae, 48(2): 523–532.

Gapon D.A. & Baena M. 2005. On the status, synonymy and tribal position of Phaeocoris Jakovlev, 1887 (Heteroptera: Pentatomidae). Zoosystematica Rossica, 14(1): 61–68. https://doi.org/10.31610/zsr/2005.14.1.61

Ghauri M.S.K. 1988. A revision of Asian species of the genus Halys Fabricius based on the type material (Insecta, Heteroptera, Pentatomidae, Pentatominae). Entomologische Abhandlungen. Staatliches Museum für Tierkunde Dresden, 51(6): 77–92.

Ghosh L.K., Biswas B., Ghosh M. & Chakraborty S.P. 1997. Hemiptera. In: State fauna series, 6. Fauna of Delhi: 215–260. Calcutta: Zoological Survey of India.

Gupta R. & Pathania P.C. 2017. Report on hemipteran pest diversity on apple plantations (Malus domestica Borkh.) in Jammu and Kashmir State of India. Records of the Zoological Survey of India, 117(4): 356–366. https://doi.org/10.26515/rzsi/v117/i4/2017/121290

Hegde V. 1995. On Heteroptera (Insecta) from the Eastern Ghats, India. Records of the Zoological Survey of India, occasional Papers, 168: 1–89.

Kandhro R., Khatri I., Rustamani M.A., Ahmed Z. & Abro N. 2016. Some stink bugs (Heteroptera: Pentatomidae) from Tando Jam Sindh. Science international (Lahore), 28(1): 187–189.

Kerzhner I.M. 2006. The etymology of Fabrician generic names in Hemiptera. Zoosystematica Rossica, 15(2): 304. https://doi.org/10.31610/zsr/2006.15.2.304

Kirkaldy G.W. 1900. On the nomenclature of the genera of the Rhynchota, Heteroptera and auchenorrhynchous Homoptera. The Entomologist, 33(449): 262–265. https://doi.org/10.5962/bhl.part.3889

Kirkaldy G.W. 1909. Catalogue of the Hemiptera (Heteroptera) with biological and anatomical references, lists of foodplants and parasites, etc, 1. Cimicidae. Berlin. xl + 392 p. https://doi.org/10.5962/bhl.title.15205

Konstantinov F.V. & Gapon D.A. 2005. On the structure of the aedeagus in shield bugs (Heteroptera, Pentatomidae): 1. Subfamilies Discocephalinae and Phyllocephalinae. Entomologicheskoe obozrenie, 84(2): 334–352 (in Russian; English translation: Entomological Review, 2005, 85(3): 221–235).

Maxwell-Lefroy H. 1909. The insect fauna of Tirhut I. – Rhynchota Heteroptera. Records of the Indian Museum, 3: 301–338. https://doi.org/10.26515/rzsi/v3/i4/1909/163293

Memon N., Ahmad I. & Perveen R. 2002. Redescription of the type species of Halys Fabricius nomen novum for Cimex dentatus Fabricius (Hemiptera: Pentatomidae: Pentatominae: Halyini), a preoccupied name of genus Halys Fabricius with a note on its independent status from Halys serrigera (Westwood). Journal of natural History and Wildlife, 1: 50–54.

Memon N., Meier R., Manan A. & Su K. F-y. 2006. On the use of DNA sequences for determining the species limits of a polymorphic new species in the stink bug genus Halys (Heteroptera: Pentatomidae) from Pakistan. Systematic Entomology, 31(4): 703–710. https://doi.org/10.1111/j.1365-3113.2006.00350.x

Memon N., Parveen R., Ahmad I. & Shaikh A.M. 2016. Description of a new species of oriental genus Halys (Pentatomoidae [sic!]: Pentominae [sic!]: Halyini) from Lahore, Punjab, Pakistan. Pakistan Journal of Zoology, 48(4): 977–980. https://doi.org/10.17582/journal.pjz/2017.49.4.1301.1306

Memon N., Parveen R., Ahmad I. & Shah N.A. 2017. Description of two new species of oriental genus Halys Fabricius (Pentatomoidae [sic!], Pentominae [sic!], Halyini) from Sindh, Pakistan with a key of its nine asian species. Pakistan Journal of Zoology, 49(4): 1301–1306. https://doi.org/10.17582/journal.pjz/2017.49.4.1301.1306

Paiva C.A. 1907. Records of Hemiptera and Hymenoptera from the Himalayas. Records of the Indian Museum, 1: 13–20. https://doi.org/10.26515/rzsi/v1/i1/1907/163419

Pajni H.R. & Sidhu C.S. 1982. A report on the pentatomid fauna of Chandigarh and surrounding areas (Heteroptera: Hemiptera). Research Bulletin of the Panjab University, Science, 33(3–4): 177–181.

Raina Surbhi T.N.K. & Kumari A. 2015. Male meiosis in two species of pentatomid bugs (Order: Hemiptera: Suborder: Heteroptera) from Jammu Region. International Journal of recent scientific Research, 6(12): 7944–7950.

Salini S. 2019. Revision of the genus Halys (Hemiptera: Pentatomidae) with description of a new species from India. Zootaxa, 4586(2): 351–375. https://doi.org/10.11646/zootaxa.4586.2.9

Schaefer C.W. 1968. The homologies of the female genitalia in the Pentatomoidea (Hemiptera–Heteroptera). Journal of the New York Entomological Society, 76: 87–91.

Scudder G.G.E. 1959. The female genitalia of the Heteroptera: morphology and bearing on classification. Transactions of the Entomological Society of London, 111(14): 405–467. https://doi.org/10.1111/j.1365-2311.1959.tb02873.x

Shaikh A.M., Memon N. & Shah M.A. 2011. A new halyine species of Halys Fabricius (Hemiptera, Pentatomidae, Pentatominae) from Miani Forest, Sindh, Pakistan. Sindh University Research Journal (Science Series), 43(2): 157–160.

Stål C. 1876. Enumeratio Hemipterorum. Bidrag till en Förteckning öfver alla hittills kända Hemiptera, Jemte Systematiska Meddelanden. Kungliga Svenska Vetenskapsakademiens Handlingar, 14(4): 1–162.

Stoll C. 1778. Natuurlyke en naar ‘t leven naauwkeurig gekleurde afbeldingen en beschryvingen der Wantzen in aller vier waerelds deelen Europa, Asia, Africa en America hishoudende, by een verzameld en beschreeven door Caspar Stoll [Représentation exactement colorée d’après nature des Punaises, qui se trouvent dans les quatre parties du monde, l’Europe, l’Asie, l’Afrique et l’Amérique; rassemblées et décrites par Caspar Stoll]. Amsterdam: Jan Christiaan Sepp. 172 p. + xli pls.

Štys P. 1961. Morphology of the abdomen and female ectodermal genitalia of the Trichophorous Heteroptera and bearing on their classification. In: Strouhal H. & Beier M. (Eds). XI. Internationaler Kongress für Entomologie, Wien, 17. bis 25. August 1960. Verhandlungen, 1: 37–43. Wien: Organisationskomittee des XI. Internationalen Kongresses für Entomologie.

Tembe S., Shouche Y. & Ghate H.V. 2014. DNA barcoding of Pentatomomorpha bugs (Hemiptera: Heteroptera) from Western Ghats of India. Meta Gene, 2: 737–745. https://doi.org/10.1016/j.mgene.2014.09.006

Tennent J.E. 1861. Sketches of the natural history of Ceylon with narratives and anecdotes illustrative of the habits and instincts of the mammalia, birds, reptiles, fishes, insects, &c. including a monograph of the elephant and a description of the modes of capturing and training it with engravings from original drawings. London: Longman, Green, Longman, and Roberts. xxv + 500 p. https://doi.org/10.5962/bhl.title.95239

Thunberg C.P. 1783. Dissertatio entomologica novas insectorum species, sistens, cujus partem secundam, cons. exper. facult. med. Upsal., publice ventilandam exhibent, 2: 29–52 + 1 pl. Upsala: Johan. Edman.

Ullah S., Ahmad H. & Rafi M.A. 2017. Taxonomy of superfamily Pentatomoidea (Hemiptera: Heteroptera) from SwathPakistan. Journal of Entomology and Zoology Studies, 5(1): 725–731.

Verhoeff C. 1893. Vergleichende Untersuchungen über die Abdominalsegmente der weiblichen Hemiptera–Heteroptera und –Homoptera, ein Beitrag zur Kenntnis der Phylogenie derselben. Verhandlungen des Naturhistorischen Vereins der Preussischen Rheinlande Westfalens und des Reg.-Bezirks Osnabruck, 50(2): 307–374. https://doi.org/10.1002/mmnd.48018930210

Walker F. 1867. Catalogue of the specimens of Hemiptera Heteroptera in the collection of the British Museum, 1. London: E. Newman. 240 p. https://doi.org/10.5962/bhl.title.118688

Volume, Year: