A review of the cytogenetically studied Odonata species of North India

Rec. zool. Surv. India: Vol. 124(3)/273-286, 2024 DOI: 10.26515/rzsi/v124/i3/2024/170418 ISSN (Online) : 2581-8686 ISSN (Print) : 0375-1511 * Author for correspondence Introduction Dragonflies comprise three suborders - Anisoptera (Dragonflies), Zygoptera (Damselflies) and Anisozygoptera (Subramanian et al., 2024). Worldwide, 686 odonate genera with 6407 species are known, while 504 species of 157 genera are present in India (Paulson et al., 2024; Subramanian et al., 2024). Odonates, due to their close association with the wetlands respond rapidly to anthropogenic or natural alterations in their habitat (Corbet, 1999; Silva et al., 2010; Oliveira Júnior et al., 2015). Odonates are used as model organisms to study behavioural ecology, genetics, and evolutionary biology (Cordoba-Aguilar, 2008). There are many reports about tolerance to physicochemical factors in Odonates (Carchini & Rota, 1985) accumulation of metals in larvae, effects of increased water temperature, sewage discharge and pesticide pollution (Thorp & Diggins, 1982; Watson et al., 1982). Cytologically, the order Odonata is well studied and it plays an important role in describing the evolutionary Abstract North India’s diverse geography and climate make it the most biodiversity-rich region in the country. Various species (flora and fauna) have been studied cytogenetically to study genetic diversity and their evolutionary relationships. The present study aims to compile the cytogenetic data of Odonata species available in the region. A total of 90 cytogenetically studied Odonata species have been listed. The chromosome number varies from n=12 to n=14 in all the studied species. Sex determination is XX/X0 type except for Nychogomphus duaricus (Fraser, 1964) and Scalmogomphus schmidti (Fraser, 1937). m chromosomes are present in 84% of the studied species. Morphological variations in the different populations of species or chromosome markers have also been enlisted. This will provide a holistic view of the number of species studied in a particular state/UT in North India and evaluate the genetic variations/ adaptations in the geographically isolated Odonata. The review also identifies gaps in the existing literature and highlights the need for further research on the cytogenetics of species in North India. Keywords:Keywords: Cytogenetic, North India, Odonata, Variations A review of the cytogenetically studied Odonata species of North India Hardeep Singh and Gurinder Kaur Walia* Department of Zoology and Environmental Sciences, Punjabi University, Patiala – 147002, Punjab, India; E-mail: gurinderk[email protected].in relationships between different groups of Odonata (Dasgupta, 1957). In Odonata, the centromere is of diffused type which is responsible for a large number of variations either due to chromosomal fusion/fragmentation or the presence/absence of m-chromosomes in the populations (Kuzenstova and Golub, 2020). The ancestral chromosome number is n=9, which increased in stepwise manners as specialization increases in the order (Kiauta, 1967) m chromosomes are the smallest or equal to the half size of immediately large chromosome and show negative heterozygosis (Mola, 1992). There are two different hypotheses regarding the origin of m chromosomesgradual diminution in the volume of normal autosomes until they eventually disappeared (Oguma, 1930; Dasgupta, 1957; Cumming, 1964) and the normal fragmentation of autosomes (Kiauta, 1968). The fragmentation can occur at any time, at any place which is responsible for variations in the size of m chromosomes in different and in the same specimen. Fusions and fragmentations play an important role in the variations in Vol 124(3) | 2024 | www.recordsofzsi.com Zoological Survey of India 274 A review of the cytogenetically studied Odonata species of North India the odonates chromosomes and these variations persist over generations through the action of natural selection and also play a role in speciation (Bueno, 1982). Northern India is one of the most climatically diverse regions where the temperature ranges from 35°C to 50 °C during summer (Thar desert, Rajasthan, Delhi) and dips to zero temperature in the winter months. Snowfall occurs in Himachal Pradesh, Ladakh, Jammu and Kashmir and Uttarakhand (Rowell, 1980; Kaul, 1998; Singh, 2003). The Indian Monsoon is responsible for the heavy rainfall as it brings moisture northwards from the Indian Ocean (Datta & Gupta, 1968; Dimri, 2004; Wang, 2006). It is characterized by the diverse range of forests (Tropical dry deciduous forests, subtropical broadleaved hill forests, Himalayan moist temperate forests, Himalayan dry temperate forests and Sub-alpine forests) (Roy et al., 2015) and wetlands which vary in topography, climate and water quality (Prasad et al., 2002). It provided suitable habitat for numerous flora and fauna. A total, of 2356 insect species are present in Himachal Pradesh, 1941 species in Uttar Pradesh, 1049 species in Rajasthan, 986 species in Delhi, 695 species in Jammu and Kashmir and 557 species in Haryana (Chandra, 2011). The Odonata species are highest in Uttrakhand (127) and decrease subsequently in Himachal Pradesh (122), Uttar Pradesh (70), Jammu and Kashmir (67), Rajasthan (51), Delhi (48), Punjab (41), Haryana (26) and Chandigarh (41) (Table 1, Subramanian et al., 2024). The present study involves Table 1. States/UT of North India with their Odonata species counts (Subramanian et al., 2024) and cytogenetically studied species S. No. States/UT Number of Odonata species No. of species studied for cytogenetics 1 Punjab 41 31 2 Himachal Pradesh 122 59 3 Uttarakhand 127 27 4 Jammu and Kashmir 67 10 5 Haryana 26 - 6 Rajasthan 51 - 7 Uttar Pradesh 70 3 8 Chandigarh 41 2 9 Delhi 48 - a comprehensive literature review on the cytogenetic studied species of North India, India (Table 2) and was necessary to fill the knowledge gap in the field. It provides an understanding of genetic diversity and the important genetic markers of the species in this area. This study can also boost conservation efforts and future research fields. Concluding Remarks and Future Directions In North India, 92 Odonata species have been studied cytogenetically which includes 76% of total odonate species present in Punjab, 48% of total species present in Himachal Pradesh, 21% of total species present in Uttrakhand, 15% of total species present in Jammu and Kashmir, 5% of total species present in Chandigarh and only 4% of total species present in Uttar Pradesh (Figure 1). The chromosome number is n=12 in the family Gomphidae, n=13-14 in the family Aeshnidae, n=13 in the family Macromiidae, n=13-12 in the family Libellulidae, n=13 in the family Cordulogastridae, Chlorogomhidae, Platycnemididae and Synlestidae, n=14 in family Coenagrionidae. m chromosome is present in 90% of studied Odonata species. However, certain species populations show variations in the chromosome complement like the presence/absence of the m chromosome, increase/decrease in the Vol 124(3) | 2024 | www.recordsofzsi.com Zoological Survey of India 275 Singh and Walia Table 2. List of Cytogenetically studied species of North India S.No. Species name Location Chromosome number Reference Remarks ANISOPTERA a) Family: Gomphidae 1Anisogomphus bivittatus (Selys, 1854) Dehradun (U.K.) n=12m Tyagi, 1982 - Andretta (H.P.) Walia & Chahal 2020 - 2 Anisogomphus occipitalis (Selys, 1854) Dehradun (U.K.) n=12m Tyagi, 1978b, 1982 - 3 Burmagomphus arboreus Lieftinck, 1964 Dehradun (U.K.) n=12m Tyagi, 1978b, 1982 - 4 Burmagomphus divericatus Lieftinck, 1964 Badhaghat and Andhretta (H.P.) n=12m Walia et al., 2021 - 5 Burmagomphus pyramidalis Laidlaw, 1922 Dehradun (U.K.) n=12m Tyagi 1977, 1982 4 larger, 4 medium-sized and 2 smaller chromosomes observed Dadhahur (Pb.) n=12m Chahal, 2019 - 6 Burmagomphus sivalikensis Laidlaw, 1922 Andhretta (H.P.) n=12m Walia et al., 2021 - 7 Burmagomphus williamsoni Forster, 1914 Dadhahur (Pb.) n=12m Chahal, 2013 - 8 Davidius zallorensis Hagen in Selys, 1878 Mansar and Surinsar Lake, Jammu (J.K.) n=12m Chahal, 2019 - 9 Ictinogomphus angulosus (Selys, 1854) Mansar and Surinsar Lake, Jammu (J.K.) n=12m Chahal, 2019 - 10 Ictinogomphus rapax (Rambur, 1842) Dehradun (U.K.) n=12m Tyagi, 1982 - Renukaji and Shah Talai (H.P.) Chahal, 2019 - 11 Lamelligomphus biforceps (Selys, 1878) Andhretta (H.P.) n=12m Chahal, 2019 - 12 Nepogomphus modestus (Selys, 1878) Palampur and Andhretta (H.P.) n=12m Waliaet al. 2006 Largest X chromosome Palampur and Andhretta (H.P.) Guchupani (U.K.) Walia & Chahal, 2014 X is medium sized Vol 124(3) | 2024 | www.recordsofzsi.com Zoological Survey of India 276 A review of the cytogenetically studied Odonata species of North India 13 Nychogomphus duraricus (Fraser, 1964) Dehradhun (U.K.) n=12, 13m Tyagi, 1977, 1985 neoXY Lachhiwala (U.K.) n=12m Chahal, 2019 - 14 Onychogomphus grammicus (Rambur, 1842) Chamarheri and Harike Wetland (Pb.) n=12m Chahal, 2019 - 15 Scalmogomphus schmidti (Fraser, 1937) Dehradhun (U.K.) n=12m, neo XY Tyagi, 1978b, 1985 - 16 Paragomphus lineatus (Selys, 1850) Dehradun (U.K.) n=12m Tyagi, 1982 - Barnala (Pb.) Walia & Chahal, 2014 Largest X chromosome 17 Platygomphus dolabratus Selys, 1854 Dadhahur and Harike Wetland (Pb.) n=12m Chahal, 2019 - 18 Scalmogomphus bistrigatus (Hagen and Selys, 1854) Dehradun (U.K.) n=12m Tyagi, 1978b, 1985 - Lacchiwala (U.K.), Andretta (H.P.) n=12m Chahal, 2019 Family: Aeshnidae 19 Anaciaeschna jaspidea (Burmeister, 1839) Khajjiar (H.P.) n=13m Walia & Sandhu (1999) - 20 Anax immaculifrons Rambur, 1842 Kandaghat (H.P.) n=14m Walia et al. 2018 - 21 Anax nigrofasciatus Fraser, 1935 Khajjiar (H.P.) n=13m Sandhu & Malhotra (1994) - Khajjiar (H.P.) n=14m Walia & Sandhu (1999) - Kandaghat (H.P.) n=14m Walia et al.(2018) - 22 Anax parthenope (Selys, 1839) Jammu & Kashmir, Himachal Pradesh n=13m Sandhu & Malhotra (1994) - 23 Cephalaeschna sp. Selys, 1883 Khajjiar (H.P.) n=13m Sandhu & Malhotra (1994) - 24 Gynacantha eschna sikkima (Karsch, 1891) Dalhousie (H.P.) n=13m Walia et al. (2016) One largest autosomal bivalent present 25 Gynacantha hyaline Selys, 1882 Dehradun (U.K.) n=14m Tyagi 1978a,b - 26 Gynacantha subinterrupta Rambur, 1842 Andretta, Palampur (H.P.) n=14m Walia & Somal, 2020 One large autosomal bivalent Vol 124(3) | 2024 | www.recordsofzsi.com Zoological Survey of India 277 Singh and Walia Family: Macromidae 27 Macromia moorei Selys, 1874 Andretta (H.P.) n=13m Walia & Chahal (2018) Family: Cordulgastridae 28 Cordulegaster brevistigma Selys, 1854 Summer Hills, Shimla (H.P.) n=13m Walia & Chahal, 2019 X chromosome is the Second largest from the compliment Family: Chlorogomphidae 29 Watanabeo petalia atkinsoni (Selys, 1878) Summer Hills, Shimla (H.P.) n=13m Walia & Chahal, 2019 Largest X chromosome Family: Libellulidae 30 Aciosoma panorpoides Rambur, 1842 Jammu and Kashmir, U.P. n=13m Walia & Sandhu, 1998 - Harike Wetland (Pb.) n=12m Kaur, 2016 Reduction in chromosome number due to the absence of the m chromosome; Largest and bipartite X chromosome 31 Brachydiplax sobrina (Rambur, 1842) Harike Wetland (Pb.) n=13m Kaur, 2016 - 32 Brachythemis contaminata Fabricius, 1793 Chandigarh n=13m Mittal & Gandhi, 1982 The largest autosomal bivalent present Punjab, Himachal Pradesh n=13m Walia and Sandhu, 1998 Largest X chromosome Punjab, Himachal Pradesh n=13m Walia et al., 2010 The largest autosomal bivalent present Anandpur Sahib, Harike Wetl and (Pb.) n=13m Kaur, 2016 - 33 Bradinopyga geminata Rambur, 1842 Barnala (Pb.) n=13m Kaur, 2016 - 34 Crocothemis servilia Drury, 1773 Chandigarh n=13m Mittal & Gandhi, 1982 One large autosomal bivalent present Punjab, U.P. Walia & Sandhu, 1998 Largest X chromosome Punjab, Himachal Pradesh Walia et al., 2010 - Bahadurgarh (Pb.) Kaur, 2016 - Vol 124(3) | 2024 | www.recordsofzsi.com Zoological Survey of India 278 A review of the cytogenetically studied Odonata species of North India 35 Diplacodes trivialis Rambur, 1842 Patiala (Pb.) n=13m Kaur, 2016 - 36 Neurothemis fulvia (Drury, 1773) Paonta Sahib (H.P.) n=13m Kaur, 2016 - 37 Neurothemis tullia Drury, 1773 Harike Wetland (Pb.) n=13m Kaur, 2016 - 38 Orthetrum anceps Schneider, 1845 Dalhousie (H.P.) n=13m Kaur, 2016 - 39 Orthetrum cancellatum Pathankot (Punjab) n=13m Dasgupta, 1957 - 40 Orthetrum glaucum Brauer, 1865 Andhretta (H.P.) n=13m Kaur, 2016 - 41 Orthetrum pruinosum Neglectum (Rambur, 1842) Punjab, Himachal Pradesh n=13m Walia et al., 2010 - Patiala (Pb.) Kaur, 2016 - 42 Orthetrum sabina Drury, 1773 Punjab, Himachal Pradesh n=13m Walia et al., 2010 - Anandpur Sahib (Pb.) Kaur, 2016 - 43 Orthetrum taeniolatum Schneider, 1845 Punjab, Jammu & Kashmir n=11m Walia et al., 2014 Two autosomal bivalent are extralarge due to fusions and show interstitial chiasmata Dalhousie (H.P.) n=13, 12m Kaur, 2016 In n=12, 3 autosomal bivalents are large in size 44 Orthetrum triangulare (Selys, 1878) Andhretta, Giripul, Bilaspur (H.P.), Nainital (U.K.) n=13m Kaur, 2016 - 45 Palpopleura sexmaculata Fabricius, 1787 Giripul, Andhretta (H.P.) n=13m Kaur, 2016 Largest X chromosome 46 Pantala flavescens Fabricius, 1798 Jammu & Kashmir and Punjab n=13m Walia & Sandhu, 1998 - Surrounding area of Harike Wetland (Pb.) n=12m Walia et al., 2011 The largest autosomal bivalent present due to the fusion of chromosomes Patiala, Harike Wetland (Pb) n=13m Kaur, 2016 - Vol 124(3) | 2024 | www.recordsofzsi.com Zoological Survey of India 279 Singh and Walia 47 Potamarcha congener (Rambur, 1842) Nainital (H.P.) n=12m Sandhu & Walia, 1995 - 48 Rhyothemis variegata (Linnaeus, 1763) Harike Wetland (Pb.) n=13m Kaur, 2016 - 49 Tholymis tillarga Fabricius, 1798 Dehradun (U.K.) n=13, 12m Kaur, 2016 2 types of cells: 1 with mchromosomes and other without m chromosomes 50 Trithemis aurora Burmiester, 1839 Punjab, Himachal Pradesh n=13m Walia et al., 2010 - Bilaspur, Andhretta (H.P.) Kaur, 2016 - 51 Trithemis festiva Rambur, 1842 Giripul, Andhretta, Bilaspur (H.P.) n=12m Kaur, 2016 Reduction in chromosome number due to the absence of m chromosomes 52 Trithemis pallidinervis Kirby, 1889 Nainital (U.K.) n=13m Kaur, 2016 - 53 Urothemis signata Rambur, 1842 Harike Wetland (Pb.) n=13m Kaur, 2016 - 54 Zygonyx torridus (Kirby, 1889) Giripul, Andhretta, Dalhousie (H.P.) n=13m Kaur, 2016 - 55 Zyxomma petiolatum Rambur, 1842 Dehradun (U.K.) n=13m Kaur, 2016 - Zygoptera Family: Calopterygidae 56 Neurobasis chinensis (Linnaeus, 1758) Narkanda, Bhunar (H.P.) n=13m Walia & Sandhu, 2002 Largest X chromosome Andretta, Mcleodganj (H.P.) n=13m Walia et al. 2016 Largest autosomal bivalent can be easily identified. Solan, Bilaspur (H.P.), Lachhiwala (U.K.) n=12m Walia & Katnoria, 2018 variation seen in the chromosome size of specimens from 3 populations Vol 124(3) | 2024 | www.recordsofzsi.com Zoological Survey of India 280 A review of the cytogenetically studied Odonata species of North India Family: Chlorocyphidae 57 Aristocyphatri fasciata (Selys, 1853) Andretta, Kullu, Chamba (H.P.) n=12m Katnoria (2020) The x chromosome is the second largest and shows bipartite behaviour 58 Aristocypha quadrimaculata (Selys, 1853) Andreta, Kullu, Chamba (H.P.) n=12m Katnoria (2020) One large autosomal bivalent 59 Libellago lineata (Burmeister, 1839) Himachal Pradesh n=13,12m Walia et al., 2018 Morphological variations in the chromosomes of both complements 60 Paracyphauni maculata (Selys, 1853) Andretta, Kullu, Chamba (H.P.) n=12m Katnoria (2020) One large autosomal bivalent Family: Euphaeidae 61 Anisopleura lestides Selys, 1853 Andretta (H.P) Dehradun (U.K.) Udampur (Jammu) n=13, 12m Walia & Katnoria, 2017 Reduction in chromosome number occurs due to the absence of m chromosome 62 Bayadera indica (Selys, 1853) Andretta, Kullu, (H.P.), Dehradun (U.K.) n=13m Walia & Katnoria, 2017 - Family: Platycnemididae 63 Subfamily-Calicnemia Calicnemia eximia (Selys, 1863) Andreta, Palampur, Chamba (H.P.) n=13m Devi, 2020 - 64 Calicnemia chromothorax (Selys, 1891) Andretta (H.P.) n=13m Walia & Devi, 2020 Bipartite X chromosome 65 Calicnemia renifera (Selys, 1896) Bilaspur (H.P.) n=13m Walia & Devi, 2020 - 66 Subfamily Platycnemidinae Copera annulata (Selys, 1863) Mansar (J&K) n=13m Walia, 2012 - 67 Copera marginipies (Rambur, 1842) Andretta, Una, Bilaspur (H.P.) Mansar lake (Jammu) n=13m Walia & Devi, 2018 - Vol 124(3) | 2024 | www.recordsofzsi.com Zoological Survey of India 281 Singh and Walia 68 Copera vittata assamensis Laidlaw, 1914 Udampur (J&K) n=13m Walia, 2012 - Renuka lake, Sirmour (H.P.) Walia & Devi, 2018 - 69 Subfamily: Disparoneurinae Prodasineura nigra Fraser, 1922 Bilaspur (H.P.) n=13m Devi, 2020 - Family: Coenagrrionidae 70 Aciagrion hisopa (Selys, 1876) Dalhousie (H.P.) n=14m Sandhu & Walia, 1993 - Andretta, Bilaspur (H.P.) Nainital (U.K.) Kaur, 2018 X chromosome Showed bipartite behaviour 71 Aciagrion palladium Selys, 1891 Bilaspur (H.P.) n=14m Kaur, 2018 - 72 Agriocnemis femina (Brauer, 1868) Patiala, Harike wetland, Muktsar (Pb.) n=14m Walia & Hallen, 2016a - Kaur, 2018 73 Agriocnemis pygmea (Rambur, 1842) Patiala, Harike wetland, Muktsar (Pb.) Dehradun (U.K.) Bilaspur (H.P.) n=14m Kaur, 2018 - 74 Agriocnemis parvum (Selys, 1876) Bilaspur (H.P.) n=14m Kaur, 2018 - 75 Ceriagrion cerinorubellum (Brauer, 1865) Harike wetland, Muktsar (Pb.) Renukaji (U.K.) n=14m Walia & Hallen, 2016a X chromosome Showed bipartite behaviour Kaur, 2018 76 Ceriagrion coromandelianum (Fabricius, 1798) Harike wetland, Muktsar (Pb.) Dehradun (U.K.) Kaur, 2018 - Rajpura (Pb.) n=14m Sandhu & Walia, 1994 - Chamba, Narkkanda, Nahan, Khajjiar (H.P.) Sandhu et al., 1994 Largest X chromosome Jammu &Kashrmir, Punjab, Himachal Pradesh, UP n=14m, 30-50 Walia & Sandhu, 1999 Fragmentation of chromosomes due to pollution