CIBTech Journal of Zoology ISSN: 2319–3883

Online, International Journal, Available at [Link]

2024 Vol.13, pp.360-374/Rajendra
Review Article (Open Access)

BIODIVERSITY OF PREDATORS AND PARASITOIDS OF

CALAPHIDINE APHIDS (HEMIPTERA: APHIDIDAE) AND
THEIR HOST PLANT ASSOCIATIONS IN INDIA

*Rajendra Singh
Department of Zoology, Deen Dayal Upadhyaya Gorakhpur University,
Gorakhpur, U.P., India

*Author for Correspondence: rsinghgpu@[Link]

ABSTRACT
The present article lists the predators and parasitoids of the calaphidine aphids (Calaphidinae: Aphididae:
Hemiptera) infesting different food plants and their distribution in various states and union territories of
India. These predators belong to four orders of the class Insecta: Coleoptera (Coccinellidae), Diptera
(Syrphidae), Hemiptera (Anthocoridae, Geocoridae) and Neuroptera (Chrysopidae, Coniopterygidae,
Hemerobiidae). A total of 48 species of predators from various taxa were recorded to feed 17 species of the
calaphidine aphids on 13 food plant species across 16 states/union territories of India. The highest number
of predators belonged to the families Coccinellidae (32 species) and Syrphidae (5 species). Most of the
tritrophic associations (triplets, predators-preys-host plants) of these predators were reported in Manipur
(36 triplets), Jammu & Kashmir (20 triplets), Uttarakhand (13 triplets) and other states/union territories
with fewer than 10 triplets. All the parasitoids of calaphidine aphids belong to a single subfamily
Aphelininae (Aphelinidae: Hymenoptera) and Aphidiinae (Braconidae: Hymenoptera) in India. A total of
15 species of parasitoids were observed parasitising calaphidine aphids infesting on 11 host plants in 7
states/union territory of India. Most of the tritrophic associations (triplets, parasitoids-hosts-host plants) of
these parasitoids were reported from Jammu & Kashmir (20 triplets), followed by Meghalaya (5 triplets),
Uttarakhand (3 triplets), Himachal Pradesh and West Bengal (each of 2 triplets), Manipur and Uttar Pradesh
(single triplet).

Keywords: Aphidiinae, biocontrol, Braconidae, Chrysopidae, Coccinellidae, parasitoids, calaphidine

aphids, predators, Syrphidae.

INTRODUCTION
The aphids exclusively belong to the family Aphididae Latreille, 1802 (Aphidoidea: Hemiptera: Insecta)
which comprises 24 subfamilies (Faveret, 2024). The Calaphidinae Oestlund, 1919 the second largest
subfamily in the Aphididae. Earlier, it was treated as a tribe under the subfamily Drepanosiphinae Herrich-
Schaeffer, 1857 (Singh and Singh, 2017). The Calaphidinae, consists of 8 tribes, 81 genera and 434 species
(Favret, 2024). Calaphidine aphids are monoecious, holocyclic (sexual generation alternates with
parthenogenetic reproduction) and oligophagous. All viviparous morphs of most of the calaphidines are
alate (winged). The sexual females are usually apterous while males are mostly alate (Singh and Ghosh,
2012). In India, 50 species are recorded under 24 genera associated with 70 plant species in 30 families,
mostly trees of Betulaceae (birches), Fagaceae (beeches), Ulmaceae (elms), Anacardiaceae (cashews), and
Juglandaceae (walnuts) and herbaceous plants like Fabaceae, Poaceae and Rosaceae (Singh and Singh,
2019; Singh et al., 2023). Several species of aphids of this subfamily are economically important pests,
causing injury and transmitting viral diseases to cultivated plants such as leguminous crops, fruit, and

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Review Article (Open Access)

landscape trees. Most of the host plants of these aphids are distributed in the western to eastern Himalayas
region.
Aphid predators and parasitoids play an important role in the natural/biological control of aphids (Singh,
2001). Records of natural enemies of these aphids begin with Ghorpade (1973) who recorded a hover fly,
Allograpta javana (Wiedemann, 1824), as a predator of Therioaphis trifolii (Monell, 1882) on alfalfa
(Medicago stiva L.) from Karnataka. After a year, Pal (1974) recorded a geocorid predatory bug, Geocoris
jucundus (Fieber, 1861) feeding the same species of aphid on alfalfa from Rajasthan. Similarly, Starý and
Ghosh (1975, 1978) were the first to record parasitoids of these aphids from Meghalaya. Thereafter, a
considerable number of predators and parasitoids were recorded from several states/union territories of
India on the calaphidine aphids on several host plants of economic importance. However, these
informations are sporadic records of species, isolated studies in the life cycles of individual species, and
discrete information on the biology and ecology of a few species. Several records of natural enemies,
particularly, predators may represent erroneous records or misuse of valid taxonomic names (Agarwala and
Ghosh, 1988). The objective of this article is to enlist the natural enemies of calaphidine aphids along with
their host plants in different states/union territories of India as such checklist provides an invaluable
reference for taxonomists, researchers, academicians, conservation managers, and policymakers to their
proper use in natural/biological control against these pests.
The present checklist is based on the primary data of published literature on predators and parasitoids, e.g.
books, book chapters, journals, proceedings of conferences, and a few authentic theses available on
Shodhganga ([Link] up to September 30, 2024. The records of predators and
parasitoids of the aphids without naming their specific prey species are excluded from this checklist. In
most of the recent - past literature, there are several errors in the scientific names of the insects both
predators/parasitoids as well as their preys/hosts (aphids) and their food plants because of their modified
status and other nomenclatural decisions and clarification. The names of aphids, as well as plants that were
misspelt in the original records have been corrected where we logically ascertain the intended species. In
the present checklist, attempts have been made to provide the valid scientific names of the predators
following GBIF (2024), WSC (2024), aphids following Favret (2024), and of the plants, following (WFO,
2024). For detailed synonymy of the valid species, the above references should be consulted.

RESULTS AND DISCUSSION

A. Predators of calaphidine aphids
The data displayed in Table 1 demonstrated that a total of 48 species of predators are recorded to prey on
17 species of calaphidine aphids out of 50 species recorded (Singh et al., 2023). Predators of these species
belong to one order of class Arachnida, the order Araneae (spiders) (3 families) and four orders of class
Insecta: Coleoptera (family Coccinellidae), Diptera (family Syrphidae), Hemiptera (families Anthocoridae,
Geocoridae) and Neuroptera (Chrysopidae, Hemerobiidae). The total number of these predator are species
distributed in 16 states/union territories of India. The present checklist shows that the maximum number of
predator species are recorded in Manipur (23 species, 36 triplets) followed by Jammu & Kashmir (11
species, 20 triplets), Uttarakhand (10 species, 13 triplets) and less than 10 species in rest of 13 states of
India. Among the calaphidine aphids, Tuberculatus paiki Hille Ris Lambers, Tuberculatus indicus Ghosh,
Chromaphis juglandicola (Kaltenbach) and Taoia indica (Ghosh and Raychaudhuri) attract 14, 13, 11, 11
and 10 species of predators, respectively (Table 2).

Table 1. Number of species of predators belonging to different taxa preying on calaphidine aphids
infesting different number of host plants distributed in different states/union territories of India.

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Orders Families of Number of

predators Predator/ Prey/host Host Triplet States/
Class

parasitoid species plant s union

species species territories
Predators
Linyphiidae 1 1 1 1 1
Arachnida

Theridiidae 1 1 1 1 1
Araneae Thomisidae 1 1 1 1 1
Sub total 3 3 3 3 3
Coleoptera Coccinellidae 32 13 9 72 9
Diptera Syrphidae 5 5 5 9 8
Anthocoridae 1 1 1 1 1
Hemiptera Geocoridae 1 1 1 1 1
Sub total 2 2 2 2 2
Chrysopidae 4 5 6 8 3
Coniopterygidae 1 1 1 1 1
Insecta

Neuroptera
Hemerobiidae 2 2 2 2 2
Sub total 7 6 6 11 4
Total 48 17 13 96 16
Parasitoids
Aphelinidae 1 1 1 1 1
Insecta

Hymenoptera Braconidae 15 11 9 15 7
Total 16 12 3 3 3

1. Class: Arachnida, Order: Araneae

The order Araneae comprises spiders which are an extremely precious element of the earth’s ecosystem
being predatory devouring mostly insects and keeping their population under check (Nyffeler and
Birkhofer, 2017). But, unlike insect predators, their potential as biocontrol agents is not exploited to its
fullest. Recently, Singh et al. (2024a) reported 77 species of aphidophagous spiders in India preying on 53
species of aphids infesting 58 species of plants. Table 1 reveals that only 3 species of spiders belonging to
3 families are known to prey on different species of calaphidine aphids in India in 3 states as mentioned
below. Indeed, minimal attempts have been made to record these aphidophagous spiders in India, and an
intensive and extensive survey plan is warranted to record them in unexplored areas.
Family 1: Linyphiidae
1. Linyphia sp.
● Taoia indica (Ghosh and Raychaudhuri, 1972)
- Alnus nepalensis D. Don - West Bengal (Raychaudhuri et al., 1979; Debnath, 2020)
Family 2: Theridiidae
1. Theridion sp.
● Myzocallis (Globulicaudaphis) pakistanica (Hille Ris Lambers, 1966)
- Quercus sp. - Himachal Pradesh (Das and Raychaudhuri, 1983)
Family 3: Thomisidae
1. Indoxysticus minutus (Tikader, 1960) [syn. Xysticus minutus Tikader, 1960]
● Pterocallis (Pterocallis) affinis Chakrabarti, 1988
- Corylus colurna L. - Uttarakhand (Debnath, 1991)
Table 2. Number of calaphidine aphids preyed by number of species of ladybird beetles, host plant
species and triplets (predator - prey - host plant associations) and their distribution in India.

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Aphid species Number of Number of Number of Number of

predator host plant triplets states/union
species species territories
Betacallis sikkimensis 4 1 4 1
Chromaphis hirsutustibis 2 2 2 1
Chromaphis juglandicola 11 1 11 2
Clethrobius dryobius 1 1 1 1
Indiochaitophorus furcatus 1 1 1 1
Myzocallis pakistanica 1 1 1 1
Myzocallis polychaeta 1 1 1 1
Panaphis juglandis 6 1 6 2
Pterocallis affinis 1 1 1 1
Sarucallis kahawaluokalani 2 2 2 1
Takecallis affinis affinis 1 1 1 1
Taoia indica 10 2 11 4
Therioaphis ononidis 1 1 1 1
Therioaphis sp. 11 1 11 9
Therioaphis trifolii 4 1 4 1
Tuberculatus indicus 13 1 13 2
Tuberculatus nervatus 7 1 7 1
Tuberculatus paiki 14 2 14 2
Total 48 13 96 16

2. Class: Insecta
Order 1: Coleoptera, Family: Coccinellidae
Among the Coleoptera, the aphidophagous ladybird beetles (Coccinellidae) are distributed throughout the
world. Though several species of this family are major agricultural pests, at least 260 species are
insectivorous feeding on soft insects like aphids, scale insects, mealy bugs, whiteflies etc. in India (Omkar
and Pervez, 2004). Several species of these beetles have been used in classical and applied biocontrol of
aphids and other soft insects (Kumar and Omkar, 2023). Table 1 demonstrates that 32 species of
Coccinellidae feed the 13 species of calaphidine aphids infesting 9 species of host plants distributed in only
9 states/union territories of India with 72 predator-prey-food plant associations (triplets). Most of the
species of these ladybird beetles are reported from Manipur (21 species) followed by Jammu & Kashmir (8
species), Karnataka (5 species triplets) and less than 5 species in other states/union territories. Among the
calaphidine aphids, Tuberculatus paiki Hille Ris Lambers and Tuberculatus indicus Ghosh attract 14 and
13 species of coccinellids, respectively, only in Manipur and West Bengal states of India. Detail predator-
prey-host plant records are given below.
1. Adalia tetraspilota (Hope, 1831)
● Chromaphis juglandicola (Kaltenbach, 1843)
- Juglans regia L. - Uttarakhand (Ghosh et al., 1991); Jammu & Kashmir (Khan and Shah, 2017; Gull and
Rasheed, 2024)
● Panaphis juglandis (Goeze, 1778)
- Juglans regia L. - Uttarakhand (Ghosh et al., 1991)
● Tinocallis sp.
- Ulmus sp. - Jammu & Kashmir (Khan and Shah, 2017)

2. Alloneda dodecaspilota (Hope, 1831) [syn. Aiolocaria dodecaspilota (Hope, 1831)]

● Taoia indica (Ghosh and Raychaudhuri, 1972)

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- Alnus nepalensis D. Don - Manipur (Chakrabarti et al., 2012)

● Taoia indica (Ghosh and Raychaudhuri, 1972)
- Alnus sp. - West Bengal (Ghosh and Raychaudhuri, 1982; Debnath, 2020)
● Taoia indica (Ghosh and Raychaudhuri, 1972)
- Alnus nitida (Spach) Endl. - Manipur (Singh and Singh, 1985)
● Tuberculatus (Acanthotuberculatus) indicus Ghosh, 1972
- Quercus serrata Murray - Manipur (Singh and Singh, 1985; Chakrabarti et al., 2012)
● Tuberculatus (Orientuberculoides) nervatus Chakrabarti and Raychaudhuri, 1976
- Quercus serrata Murray - Manipur (Chakrabarti et al., 2012)
3. Brumoides suturalis (Fabricius, 1798) [syn. Brumus suturalis (Fabricius, 1798)]
● Therioaphis (Pterocallidium) trifolii (Monell, 1882)
- Medicago sativa L. - Uttar Pradesh (Chaudhary and Singh, 2012)
4. Calvia championorum Booth, 1997
● Tuberculatus (Acanthotuberculatus) indicus Ghosh, 1972
- Quercus serrata Murray - Manipur (Chakrabarti et al., 2012)
5. Calvia explanata Poorani, 2014
● Chromaphis juglandicola (Kaltenbach, 1843)
- Juglans regia L. - Jammu & Kashmir (Khan and Shah, 2017; Gull and Rasheed, 2024)
● Panaphis juglandis (Goeze, 1778)
- Juglans regia L. - Jammu & Kashmir (Khan and Shah, 2017)
● Taoia indica (Ghosh and Raychaudhuri, 1972)
- Alnus nepalensis D. Don - Sikkim (Joshi and Sangma, 2015)
● Tinocallis sp.
- Ulmus sp. - Jammu & Kashmir (Khan and Shah, 2017)
6. Calvia sykesii (Crotch, 1874)
● Taoia indica (Ghosh and Raychaudhuri, 1972)
- Alnus nepalensis D. Don - Sikkim (Joshi and Sangma, 2015)
7. Cheilomenes sexmaculata (Fabricius, 1781) [syn. Menochilus sexmaculatus (Fabricius, 1781)]
● Sarucallis kahawaluokalani (Kirkaldy, 1907)
- Unknown plant - Tripura (Agarwala et al., 1987)
● Therioaphis (Pterocallidium) trifolii (Monell, 1882)
- Medicago sativa L. - Gujarat (Patel, 2015); Karnataka (Megha et al., 2015); Uttar Pradesh (Chaudhary
and Singh, 2012)
● Tuberculatus (Orientuberculoides) nervatus Chakrabarti and Raychaudhuri, 1976
- Quercus serrata Murray - Manipur (Somen Singh et al., 1995)
● Tuberculatus (Orientuberculoides) paiki Hille Ris Lambers, 1974
- Quercus serrata Murray - Manipur (Singh et al., 1985)
8. Coccinella septempunctata Linnaeus, 1758
● Chromaphis juglandicola (Kaltenbach, 1843)
- Juglans regia L. - Jammu & Kashmir (Khan and Shah, 2017; Gull and Rasheed, 2024)
● Myzocallis (Neodryomyzus) polychaeta David, 1969
- Quercus sp. - Uttarakhand (Ghosh et al., 1991)
● Panaphis juglandis (Goeze, 1778) - Juglans regia L. - Jammu & Kashmir (Khan and Shah, 2017)
● Takecallis affinis affinis Ghosh, 1986
- Bambusa sp. - Uttarakhand (Ghosh et al., 1991)
● Taoia indica (Ghosh and Raychaudhuri, 1972)
- Alnus nepalensis D. Don - Uttarakhand (Ghosh et al., 1991); West Bengal (Ghosh and Raychaudhuri,
1982; Debnath, 2020)
● Therioaphis ononidis (Kaltenbach, 1846)

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- Unknown aphid - Unknown plant (Agarwala and Ghosh, 1988)

● Therioaphis (Pterocallidium) trifolii (Monell, 1882)
- Medicago sativa L. - Uttar Pradesh (Chaudhary and Singh, 2012)
● Tuberculatus (Acanthotuberculatus) indicus Ghosh, 1972
- Quercus serrata Murray - Manipur (Chakrabarti et al., 2012)
● Tuberculatus (Orientuberculoides) paiki Hille Ris Lambers, 1974
- Quercus serrata Murray - Manipur (Singh et al., 1985); West Bengal (Chakrabarti et al., 2012)
9. Coccinella transversalis Fabricus, 1781
● Therioaphis (Pterocallidium) trifolii (Monell, 1882)
- Medicago sativa L. - Karnataka (Megha et al., 2015); Uttar Pradesh (Chaudhary and Singh, 2012)
● Tuberculatus (Acanthotuberculatus) indicus Ghosh, 1972
- Unknown plant - India (Chakrabarti et al., 2012)
● Tuberculatus (Orientuberculoides) paiki Hille Ris Lambers, 1974
- Quercus serrata Murray - Manipur (Singh et al., 1985)
10. Cryptogonus bimaculatus Kapur, 1948
● Tuberculatus (Orientuberculoides) paiki Hille Ris Lambers, 1974
- Quercus serrata Murray - Manipur (Singh et al., 1985; Chakrabarti et al., 2012)
11. Harmonia dimidiata (Fabricius, 1781)
● Chromaphis juglandicola (Kaltenbach, 1843)
- Juglans regia L. - Jammu & Kashmir (Gull and Rasheed, 2024)
● Tuberculatus (Acanthotuberculatus) indicus Ghosh, 1972
- Quercus serrata Murray - Manipur (Shantibala, 1989)
● Tuberculatus (Orientuberculoides) nervatus Chakrabarti and Raychaudhuri, 1976
- Unknown plant - Manipur (Chakrabarti et al., 2012)
● Tuberculatus (Orientuberculoides) paiki Hille Ris Lambers, 1974
- Quercus serrata Murray - Manipur (Singh et al., 1985)
12. Harmonia eucharis (Mulsant, 1853)
● Chromaphis juglandicola (Kaltenbach, 1843)
- Juglans regia L. - Jammu & Kashmir (Khan and Shah, 2017; Gull and Rasheed, 2024)
● Panaphis juglandis (Goeze, 1778)
- Juglans regia L. - Jammu & Kashmir (Khan and Shah, 2017)
● Tuberculatus (Acanthotuberculatus) indicus Ghosh, 1972
- Quercus serrata Murray - Manipur (Chakrabarti et al., 2012)
● Tuberculatus (Orientuberculoides) nervatus Chakrabarti and Raychaudhuri, 1976
- Unknown plant - Manipur (Chakrabarti et al., 2012)
● Tuberculatus (Orientuberculoides) paiki Hille Ris Lambers, 1974
- Quercus serrata Murray - Manipur (Shantibala, 1989; Singh et al., 1985)
13. Harmonia octomaculata (Fabricius, 1781)
● Therioaphis (Pterocallidium) trifolii trifolii (Monell, 1882)
- Medicago sativa L. - Karnataka (Megha et al., 2015)
● Tuberculatus (Acanthotuberculatus) indicus Ghosh, 1972
- Quercus serrata Murray - Manipur (Chakrabarti et al., 2012)
● Tuberculatus (Orientuberculoides) paiki Hille Ris Lambers, 1974
- Quercus serrata Murray - Manipur (Singh et al., 1985)
14. Hippodamia convergens Guérin - Méneville, 1842
● Chromaphis juglandicola (Kaltenbach, 1843)
- Juglans regia L. - Jammu & Kashmir (Ahmad and Wani, 2014)
● Panaphis juglandis (Goeze, 1778)
- Juglans regia L. - Jammu & Kashmir (Ahmad and Wani, 2014)

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15. Hippodamia variegata (Goeze, 1777) [syn. Adonia variegata (Goeze, 1777)]
● Chromaphis juglandicola (Kaltenbach, 1843)
- Juglans regia L. - Jammu & Kashmir (Khan and Shah, 2017)
● Therioaphis (Pterocallidium) trifolii trifolii (Monell, 1882)
- Medicago sativa L. - Karnataka (Megha et al., 2015)
● Tinocallis sp.
- Ulmus sp. - Jammu & Kashmir (Khan and Shah, 2017)
16. Micraspis discolor (Fabricius, 1798) [syn. Verania discolor (Fabricius, 1798)]
● Tuberculatus (Orientuberculoides) paiki Hille Ris Lambers, 1974
- Quercus serrata Murray - Manipur (Shantibala, 1989)
17. Micraspis vincta (Gorham, 1895)
● Tuberculatus (Orientuberculoides) paiki Hille Ris Lambers, 1974
- Quercus serrata Murray - Manipur (Singh et al., 1985)
18. Oenopia conglobata (Linnaeus, 1758)
● Chromaphis juglandicola (Kaltenbach, 1843)
- Juglans regia L. - Jammu & Kashmir (Gull and Rasheed, 2024)
19. Oenopia kirbyi Mulsant, 1850
● Tuberculatus (Acanthotuberculatus) indicus Ghosh, 1972
- Quercus serrata Murray - Manipur (Chakrabarti et al., 2012)
● Tuberculatus (Orientuberculoides) paiki Hille Ris Lambers, 1974
- Quercus serrata Murray - Manipur (Singh et al., 1985)
20. Oenopia manipurensis Devi, Singh and Singh, 1991
● Tuberculatus (Orientuberculoides) paiki Hille Ris Lambers, 1974
- Quercus serrata Murray - Manipur (Shantibala, 1989)
21. Oenopia mimica Weise, 1902
● Taoia indica (Ghosh and Raychaudhuri, 1972)
- Alnus nepalensis D. Don - Sikkim (Joshi and Sangma, 2015)
22. Oenopia quadripunctata Kapur, 1963
● Tuberculatus (Acanthotuberculatus) indicus Ghosh, 1972
- Quercus serrata Murray - Manipur (Chakrabarti et al., 2012)
● Tuberculatus (Orientuberculoides) nervatus Chakrabarti and Raychaudhuri, 1976
- Quercus serrata Murray - Manipur (Singh et al., 1993)
● Tuberculatus (Orientuberculoides) paiki Hille Ris Lambers, 1974
- Quercus serrata Murray - Manipur (Singh et al., 1985)
23. Oenopia sauzeti Mulsant, 1866
● Clethrobius dryobius Chakrabarti and Raychaudhuri, 1976
- Prunus cerasus L. - Sikkim (Chakrabarti et al., 2012)
24. Oenopia sexareata (Mulsant, 1853) [syn. Coelophora sexarcata Mulsant, 1853]
● Taoia indica (Ghosh and Raychaudhuri, 1972)
- Alnus nepalensis D. Don - West Bengal (Raychaudhuri et al., 1978; Ghosh and Raychaudhuri, 1982)
● Tuberculatus (Orientuberculoides) paiki Hille Ris Lambers, 1974
- Quercus semiserrata Roxb. - Manipur (Shantibala, 1989)
25. Phrynocaria unicolor (Fabricius, 1792) [syn. Coelophora unicolor (Fabricius, 1792)]
● Taoia indica (Ghosh and Raychaudhuri, 1972)
- Alnus sp. - Manipur (Chakrabarti et al., 2012)
● Tuberculatus (Acanthotuberculatus) indicus Ghosh, 1972
- Quercus serrata Murray - West Bengal (Chakrabarti et al., 2012)
26. Propylea dissecta (Mulsant, 1850)
● Therioaphis (Pterocallidium) trifolii trifolii (Monell, 1882)

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- Medicago sativa L. - Karnataka (Megha et al., 2015)

27. Propylea luteopustulata (Mulsant, 1850) [syn. Oenopia luteopustulata Mulsant, 1850; Pania
luteopustulata (Mulsant, 1850)]
● Tuberculatus (Acanthotuberculatus) indicus Ghosh, 1972
- Quercus serrata Murray - Manipur (Chakrabarti et al., 2012)
28. Scymnus (Pullus) giganteus Kamiya, 1961
● Tuberculatus (Acanthotuberculatus) indicus Ghosh, 1972
- Unknwon aphid - Unknown plant (Chakrabarti et al., 2012)
29. Scymnus (Pullus) hilaris Motschulsky, 1858
● Tuberculatus (Orientuberculoides) paiki Hille Ris Lambers, 1974
- Quercus serrata Murray - Manipur (Singh et al., 1985; Chakrabarti et al., 2012)
30. Scymnus (Scymnus) nubilus Mulsant, 1850
● Therioaphis (Pterocallidium) trifolii (Monell, 1882)
- Unknwin aphid - Unknown plant - (Chakrabarti et al., 2012)
31. Scymnus sp.
● Taoia indica (Ghosh and Raychaudhuri, 1972)
- Alnus nepalensis D. Don - Manipur (Agarwala et al., 1980; Chakrabarti et al., 2012)
32. Synonycha grandis (Thunberg, 1781)
● Tuberculatus (Acanthotuberculatus) indicus Ghosh, 1972
- Quercus serrata Murray - Manipur (Chakrabarti et al., 2012)

Order 2: Diptera, Family: Syrphidae

Among the Diptera, the Syrphidae, commonly known as hover flies, are the most common aphid predators
and are widely distributed throughout the world. Most of the syrphids provide dual service in nature, as an
adult they help in crop pollination (Joshi et al., 2023) and as larvae, they consume several soft-bodied
insects such as aphids, scale insects, mealy bugs, whiteflies, thrips etc.; and thus form an integral part of
natural control of these insects (Kumar and Omkar, 2023). Table 1 demonstrates that only 5 species of
hover flies feed the 5 species of calaphidine aphids infesting 5 species of host plants distributed in only 8
states/union territories of India with 9 predator-prey-food plant associations (triplets). Detail predator-prey-
host plant records are given below.
1. Allograpta javana (Wiedemann, 1824) [syn. Sphaerophoria javana Wiedemann, 1824; Syrphus
javana Wiedemann, 1824]
● Therioaphis (Pterocallidium) trifolii (Monell, 1882)
- Medicago sativa L. - Assam (Agarwala et al., 1984); Bihar (Agarwala et al., 1984); Karnataka (Ghorpadé,
1973); Meghalaya (Agarwala et al., 1984); Tamil Nadu (Agarwala et al., 1984)
2. Episyrphus balteatus (De Geer, 1776) [syn. Syrphus balteatus De Geer, 1776]
● Indiochaitophorus furcatus Verma, 1970
- Ulmus wallichiana Planch. (syn. Ulmus laevigata Royle) - Jammu Kashmir (Bhagat and Lone, 1984)
3. Eupeodes confrater (Wiedemann, 1830) [syn. Metasyrphus confrater (Wiedemann, 1930)]
● Tuberculatus (Orientuberculoides) nervatus Chakrabarti and Raychaudhuri, 1976
- Quercus serrata Murray - Manipur (Somen Singh et al., 1995)
4. Eupeodes latifasciatus (Macquart, 1829) [syn. Metasyrphus latifasciatus (Macquart, 1829)]
● Chromaphis juglandicola (Kaltenbach, 1843)
- Juglans regia L. - Jammu & Kashmir (Gull and Rasheed, 2024)
5. Ischiodon scutellaris (Fabricius, 1805) [syn. Xanthogramma scutellare (Fabricius, 1805)]
● Sarucallis kahawaluokalani (Kirkaldy, 1907)
- Unknown plant - Tripura (Agarwala et al., 1987)

Order 3: Hemiptera

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The order Hemiptera includes at least 7 families of predatory bugs preying on aphids, scale insects,
mealybugs, bugs, leaf hoppers, thrips, mites, caterpillars etc. (Chellappan and Ranjith, 2023). In India,
members of 2 families, Anthocoridae and Geocoridae, one species each, are recorded as predators
of calaphidine aphids in Uttarakhand and Rajasthan, respectively (Table 1) as mentioned below.
Family 1: Anthocoridae
1. Orius bifilarus Ghauri, 1972
● Taoia indica (Ghosh and Raychaudhuri, 1972)
- Alnus nepalensis D. DonUttarakhand (Debnath, 1991)

Family 2: Geocoridae
2. Geocoris jucundus (Fieber, 1861)
● Therioaphis (Pterocallidium) trifolii (Monell, 1882)
- Medicago sativa L. - Rajasthan (Pal, 1974)

Order 4: Neuroptera
Neuroptera includes lacewings, mantisflies, antlions and their relatives and both adults and larvae are
entomophagous. They are globally distributed and generalist predators of soft-bodied insects. Recently,
Singh et al. (2024b) enlisted 32 species of aphidophagous neuropterans from India belonging to 4 families.
However, members of only three families, Chrysopidae (green lacewings), Coniopterygidae (dustywings)
and Hemerobiidae (brown lacewings) are recorded to prey on calaphidine aphids in India. Among the
chrysopids, commonly known as green lacewings, only 4 species prey on 5 species of calaphidine aphids
feeding on 6 species of food plants distributed mostly in Uttarakhand states of India. Only a single species
of dustywing and two species of brown lacewings are in India as mentioned below.
Family 1: Chrysopidae
1. Chrysopa himalayana Ghosh, 1985
● Chromaphis hirsutustibis Kumar and Lavigne, 1970
- Juglans regia L. - Uttarakhand (Debnath et al., 1988)
2. Chrysoperla zastrowi sillemi (Esben - Petersen, 1935) [syn. Chrysopa gujaratensis Ghosh, 1976;
Chrysopa punensis Ghosh, 1976; Chrysopa sanandensis Ghosh, 1977; Chrysopa sillemi Esben -
Petersen, 1935
● Betacallis sikkimensis Basu, Ghosh and Raychaudhuri, 1974
- Betula alnoides Buch.–Ham. - Uttarakhand (Dey, 2015)
● Chromaphis juglandicola (Kaltenbach, 1843)
- Juglans regia L. - Jammu & Kashmir (Khan and Shah, 2017)
● Panaphis juglandis (Goeze, 1778)
- Juglans regia L. - Jammu & Kashmir (Khan and Shah, 2017)
● Therioaphis (Pterocallidium) trifolii trifolii (Monell, 1882)
- Medicago sativa L. - Maharashtra (Tambe and Kadam, 2015)
● Tinocallis sp.
- Ulmus sp. - Jammu & Kashmir (Khan and Shah, 2017)
3. Pseudomallada alcestes (Banks, 1911) [syn. Mallada alcestes (Banks, 1911)]
● Betacallis sikkimensis Basu, Ghosh and Raychaudhuri, 1974
- Betula alnoides Buch.–Ham. - Uttarakhand (Dey, 2015)
4. Retipenna dasyphlebia (McLachlan, 1894) [syn. Chrysopa dasyphlebia McLachlan, 1894]
● Betacallis sikkimensis Basu, Ghosh and Raychaudhuri, 1974
- Betula alnoides Buch.–Ham. - Uttarakhand (Debnath et al., 1988; Dey, 2015)
Family 2: Coniopterygidae
1. Coniocompsa indica Withycombe, 1925
● Chromaphis hirsutustibis Kumar and Lavigne, 1970

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- Berberis sp. - Uttarakhand (Dey, 2015)

Family 3: Hemerobiidae
2. Micromus timidus Hagen, 1853
● Tuberculatus (Orientuberculoides) nervatus Chakrabarti and Raychaudhuri, 1976
- Quercus serrata Murray - Manipur (Somen Singh et al., 1995)
3. Micromus sp.
● Betacallis sikkimensis Basu, Ghosh and Raychaudhuri, 1974
- Betula alnoides Buch.–Ham. - Uttarakhand (Debnath, 1991)

B. Parasitoids of calaphidine aphids

The parasitoids of the calaphidine aphids belong to two families of the order Hymenoptera: Aphelinidae
(only subfamily Aphelininae, tribe Aphelinini) and Braconidae (only subfamily Aphidiinae). A total of 16
species of parasitoids belonging to different taxa were observed to parasitise 15 species of these aphids on
11 food plant species distributed in 7 states/union territories of India (Table 1).
The Aphelinidae are a small group of Chalcidoidea (Hymenoptera) containing 32 genera and a little over a
thousand species (Hayat, 1998) and is a major source of biocontrol agents of economically important insect
pest species such as coccoids, aphids and aleyrodids (Homoptera). Aphelinidae is represented by a single
species, Aphelinus argiope Walker while

Table 3. Number of species of aphidiine parasitoids parasitising the number of host aphids
calaphidine aphids host plants infested, number of tritrophic associations and their distribution in
India
Aphid species Number of Number of host Number of Distribution in
parasitoid plants triplets states/union
species territories
1. Betacallis 1 2 2 1
querciphaga
2. Betacallis 1 1 1 1
sikkimensis
Betacallis sp. 1 1 1 1
3. Chromaphis 1 1 1 1
juglandicola
4. Myzocallis ulmifoliae 1 1 1 1
5. Panaphis juglandis 2 1 2 3
6. Saltusaphis sp. 1 1 1 1
7. Shivaphis celti 4 2 4 2
Shivaphis sp. 1 1 1 1
8. Takecallis affinis 2 1 2 2
affinis
9. Takecallis 1 1 1 2
arundinariae
10. Taoia indica 1 1 1 2
11. Therioaphis trifolii 1 1 1 1
Therioaphis sp. 1 1 1 1
12. Tuberculatus indicus 1 1 1 1
Total 15 11 15 7

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Das and Chakrabarti (2023) listed 157 aphidiine species parasitising several aphid species in India. Out of
these, only 15 species are described/recorded from India parasitising 15 species of the calaphidine aphids
(in addition, 3 aphids are identified up to genric level only) infesting 11 species of host plants (in addition,
3 unknown species) distributed only in 7 states/union territories of India. All these parasitoids parasitise
only 1-2 aphid species. The maximum number of parasitoids was recorded on Shivaphis (Shivaphis) celti
Das (4 species of braconid parasitoids) in India (Table 3). It indicates that India has very limited fauna and
biodiversity of parasitoids of the calaphidine aphids of economic importance. It demonstrates that survey
programmes should be conducted regarding the biodiversity of parasitoids of these aphids in unexplored
areas. Following checklist displays the species of aphidiine parasitoids recorded/described parasitising the
calaphidine in India.
The following is the checklist of aphid parasitoids along with their host aphids infesting various food plants
in different states/union territories of India is mentioned below.
Family 1: Aphelinidae, subfamily: Aphelininae
1. Aphelinus argiope Walker, 1839 [syn. Mesidia argiope (Walker, 1839)]
● Saltusaphis sp.
- Unknown plant - Uttar Pradesh (Hayat, 1979)
Family 2: Braconidae, subfamily: Aphidiinae
1. Aphidius matricariae Haliday, 1834
● Myzocallis (Myzocallis) ulmifoliae Shinji, 1954
- Rubia cordifolia L. - Jammu & Kashmir (Bhagat and Ahmad, 1991)
2. Betuloxys assamensis (Stary, 1975) [syn. Trioxys (Betuloxys) assamensis Stary, 1975]
● Betacallis querciphaga Basu, Ghosh and Raychaudhuri, 1974
- Quercus sp. - Meghalaya (Ghosh and Raychaudhuri, 1982)
- Unknown plant - Meghalaya (Starý and Ghosh, 1975)
3. Betuloxys takecallis (Stary, 1978) [syn. Trioxys (Betuloxys) takecallis Starý, 1978; Trioxys
takecallis Starý, 1978]
● Takecallis arundinariae (Essig, 1917)
- Bambusa sp. - Meghalaya (Raychaudhuri et al., 1979); West Bengal (Starý and Raychaudhuri, 1978)
4. Binodoxys jaii (Bhagat, 1982) [Trioxys jaii Bhagat, 1982]
● Shivaphis (Shivaphis) celti Das, 1918
- Celtis australis L. - Jammu & Kashmir (Bhagat, 1982)
5. Binodoxys manipurensis Singh PM and Singh, 1986
● Shivaphis (Shivaphis) celti Das, 1918
- Celtis tetrandra Roxb. - Manipur (Singh and Singh, 1986)
6. Binodoxys takecallis (Das and Chakrabarti, 1989)
● Takecallis affinis affinis Ghosh, 1986 [syn. Takecallis himalayensis Chakrabarti, 1988]
- Bambusa sp. - Himachal Pradesh (Chakrabarti and Debnath, 2009); Uttarakhand (Das and Chakrabarti,
1989a; Chakrabarti and Debnath, 2009)
7. Ephedrus lacertosus (Haliday, 1833)
● Takecallis affinis affinis Ghosh, 1986 [syn. Takecallis himalayensis Chakrabarti, 1988]
- Bambusa sp. - Himachal Pradesh (Chakrabarti and Debnath, 2009); Uttarakhand (Das and Chakrabarti,
1989a; Sarkar, 2022)
8. Lipolexis gracilis Forster, 1862
● Therioaphis (Pterocallidium) trifolii (Monell, 1882)
- Unknown plant - Jammu & Kashmir (Starý and Ghosh, 1983; Chakrabarti and Debnath, 2009)
9. Praon himalayensis Das and Chakrabarti, 1989
● Panaphis juglandis (Goeze, 1778)
- Juglans regia L. - Himachal Pradesh (Chakrabarti and Debnath, 2009); Uttarakhand (Das and
Chakrabarti, 1989b; Sarkar, 2022)

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10. Toxares deltiger (Haliday, 1833)

● Betacallis sikkimensis Basu, Ghosh and Raychaudhuri, 1974
- Conium maculatum L. - Jammu & Kashmir (Chakrabarti and Debnath, 2009)
● Betacallis sp.
- Conium maculatum L. - Jammu & Kashmir (Bhagat, 1983); Jammu & Kashmir (Bhagat, 1984)
11. Trioxys complanatus Quilis, 1931
● Therioaphis sp.
- Vicia sativa L. - Jammu & Kashmir (Bhagat, 1984)
12. Trioxys pallidus (Haliday, 1833)
● Chromaphis juglandicola (Kaltenbach, 1843)
- Juglans regia L. - Jammu & Kashmir (Shuja-Uddin, 1982; Ahmad and Wani, 2014)
● Panaphis juglandis (Goeze, 1778)
- Juglans regia L. - Jammu & Kashmir (Ahmad and Wani, 2014; Khan and Shah, 2017)
13. Trioxys raychaudhurii Bhagat, 1990
● Taoia indica (Ghosh and Raychaudhuri, 1972)
- Alnus nepalensis D. Don - Meghalaya (Raychaudhuri et al., 1990)
● Taoia indica (Ghosh and Raychaudhuri, 1972)
- Alnus nepalensis D. Don - West Bengal (Tamili, 1988)
14. Trioxys shivaphis Takada, 1966
● Shivaphis (Shivaphis) celti Das, 1918
- Celtis australis L. - Jammu & Kashmir (Bhagat, 1984)
15. Trioxys soporensis Shuja-Uddin, 1982
● Shivaphis (Shivaphis) celti Das, 1918
- Celtis sp. - Jammu & Kashmir (Chakrabarti and Debnath, 2009)
● Shivaphis sp.
- Celtis sp. - Jammu & Kashmir (Shuja-Uddin, 1982)
16. Trioxys sp.
● Tuberculatus (Acanthotuberculatus) indicus Ghosh, 1972
- Quercus griffithii Hook.f and Thoms. ex Miq. - Meghalaya (Starý and Ghosh, 1978)

CONCLUSION
The calaphidine aphids are highly economically important infesting different food plants of economic
importance such as beeches, cashews, walnuts etc. in India. In nature, its population is regulated by their
natural enemies. In this checklist, 48 species of predators and 15 species of parasitoids of these aphids are
listed which are distributed in 16 and 7 states/union territories of India. Indeed, very few attempts have
been made to record these natural enemies in India, and an intensive and extensive survey plan is warranted
to record them in unexplored areas.

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