116 urn:lsid:arphahub.com:pub:619a5b3a-5ec8-5ff7-b0b1-5070a7c17694 urn:lsid:zoobank.org:pub:70C65CC0-001D-487B-A05D-B86A205B9582 Contributions to Entomology CTE 0005-805X 2511-6428 Senckenberg Gesellschaft für Naturforschung 10.3897/contrib.entomol.73.e109883 109883 Research Article Trichoptera Biodiversity & Conservation Systematics Recently collected Lepidostoma species (Trichoptera, Lepidostomatidae) from India, with new records Hussain Zahid 1 Majeed Aquib 1 Ali Tabraq https://orcid.org/0000-0002-8686-391X 1 Parey Sajad H. sajadzoo@gmail.com https://orcid.org/0000-0002-2094-0812 1 Insect Systematic Research Laboratory, Department of Zoology, Baba Ghulam Shah Badshah University, Rajouri, (J&K), India Baba Ghulam Shah Badshah University Rajouri India

Corresponding author: Sajad H. Parey (sajadzoo@gmail.com)

Academic editor: Simon Vitecek

 2023 23 11 2023 73 2 201 208 BC6D9819-E821-5B85-BF7B-D2FEE9F5133B 6DDD08A5-DA0A-45AE-B828-6191E63FAF27 10412270 21 07 2023 11 11 2023 Zahid Hussain, Aquib Majeed, Tabraq Ali, Sajad H. Parey This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. http://zoobank.org/6DDD08A5-DA0A-45AE-B828-6191E63FAF27

Two new records of the genus Lepidostoma Rambur are reported from India. These include L. diespiter (Malicky & Sangpradub, 2001) and L. kamba (Mosely, 1939b) collected from Himachal Pradesh and Uttarakhand in India. With these new additions, the genus Lepidostoma Rambur is represented by 51 valid species from India. Complete redescriptions of these two species with illustrations are also provided. Also, L. sonomax (Mosely, 1939) is reported from Uttarakhand for the first time. Potential threats to these species and other freshwater biota are noted.

 Genitalia northwestern Himalayas taxonomy vector graphics Science and Engineering Research Board (SERB), Govt of India Introduction

The family Lepidostomatidae was originally described by Ulmer (1903) as a subfamily of Sericostomatidae. In India, this family is represented by three genera, Lepidostoma Rambur, Paraphlegopteryx Ulmer, Zephyropsyche Weaver. These comprise 49 species of Lepidostoma, 15 species of Paraphlegopteryx, and 1 species of Zephyropsyche in the Indian fauna.

Secondary sexual characters of the maxillary palps, antennal scapes, and forewings are highly modified in Lepidostoma. These secondary sexual characters can be used to distinguish closely related species with similar genitalia. Based on wing venation, Weaver (2002) divided the genus Lepidostoma into four branches: L. hirtum branch, L. vernale branch (monophyletic), L. podogram branch, L. ferox branch.

In India, Lepidostoma is represented only by the non-monophyletic L. hirtum and L. ferox branches. Earlier contributions to Lepidostoma in India include 16 species described by Mosely (1939, 1941, 1949a, 1949b, 1949c); 5 species by Martynov (1936); 4 species by Malicky (1979, 1994, 2003, 2007), 3 species by McLachlan (1871, 1878), 2 species by Ulmer (1905, 1906), 2 species by Weaver (1989, 2002), 10 species by Parey et al. (2012, 2013, 2016, 2019), 3 species by Saini et al. (2011), and 1 species each by Navàs (1932), Ito (1964), Kimmins (1986), and Dinakaran et al. (2013), making a total of 49 species. Two additional species of Lepidostoma reported here increase the count to 51 species of this genus reported from India.

 Methods and materials

An extensive survey was conducted during 2018–2022 in the northwestern Himalayas of India. During the survey, we used a 15-watt black light tube (BioQuip, Inc) gifted by Dr. Karl M. Kjer to Sajad H. Parey during the XV International Symposium on Trichoptera hosted at Rutgers University, New Brunswick, New Jersey, USA. This light trap was powered by a sealed rechargeable 12-volt battery placed over a white sheet for 1–4 hours beginning at dusk (Fig. 3B). The trap was placed near freshwater streams, lentic habitats, and other water bodies. We took photographs of the habitats with Xiaomi Redmi Note 10S mobile camera (Figs 3A, 4A–C). The collected specimens were killed and preserved in 96% alcohol.

These collected specimens were identified to families based on their morphological characters by using the illustrated key for adults by Saini et al. (2001). To identify the specimens to species, we dissected the genitalia using fine-tipped forceps, macerated them by using the lactic acid method of Blahnik et al. (2007), then put them in 70% alcohol with a drop of glycerin for better visualization under a stereo-zoom microscope. Templates for the drawings were prepared using the Xiaomi Redmi Note 10S mobile camera, final vector graphics were prepared by using Adobe Illustrator 2020; plates of illustrations were assembled using Photoshop.

 Results Animalia Trichoptera Lepidostomatidae 5128E579-E79A-540D-B2BD-A2ED4355F920 Lepidostoma diespiter (Malicky & Sangpradub, 2001) Dinarthrum diespiter Malicky & Sangpradub, 2001; 13, figs. Description.

Adult ♂ (Fig. 1D, 1E): Scapes each with single basal process, broad basally and apically tapering to forms finger-like structure. Maxillary palps each single-segmented, medially with long projection apically somewhat tapering, covered with dense setae (Fig. 1D). Forewings each with apical forks I and II and with three post-cubital cells (Fig. 1E).

10.3897/contrib.entomol.73.e109883.figure1 C794607A-B63D-57A1-B1A2-C2FDE3976B32

Lepidostoma diespiter (Malicky & Sangpradub, 2001). A–C. Male genitalia: A. Dorsal; B. Ventral; C. Left lateral. D. Head with scape; E. Forewing; F. Phallic apparatus. AD = apicodorsal process of inferior appendage (paired); AV = apicoventral process of inferior appendage (paired); BDP= Basodorsal process of inferior appendage (paired); BP = two basal processes of antennal scape (paired); IFA = inferior appendage (paired); IX seg. = segment; PA = phallic paramere (paired); PH = phallus; X Seg = segment X.

https://binary.pensoft.net/fig/941755

Genitalia ♂ (Fig. 1A–C, F): Segment IX posterodorsally produced into triangle with round apex, segment IX dorsolateral projections rounded. Segment X in dorsal view (Fig. 1A) anterodorsally wide, constricted at mid-length then tapering towards apex, apex with small notch equipped with triangular projection; in lateral view (Fig. 1C), posterolaterally divided into two apically round lobes, dorsal lobe projected posterodorsad and pair of ventrolateral lobes projected posteroventrad, small acute triangular projection visible below ventrolateral lobes of tergum X. Inferior appendages each single-segmented, in lateral view (Fig. 1C) basal 2/3 tubular, distal 1/3 with two processes: apicodorsal process half as thick as basal 2/3 and apically truncate, apicoventral process digitate; in ventral view (Fig. 1B) each inferior appendage curved caudad, apically bilobed with clavate apicodorsal lobe twice as long as clavate apicoventral lobe. Phallus tongue-shaped in ventral view (Fig. 1B) and in lateral view (Fig. 1F) phallus apically rounded triangular with parameres placed diagonally across genitalia); basodorsal process present, serrated, and finger-like.

Female. Unknown.

 Holotype repository.

In the collection of Hans Malicky, Lunz am See, Austria.

 Material examined.

India (new record): Himachal Pradesh: Andretta, 32°02'22.2"N, 76°35'16.5"E, 31-v-2022, 962 m, 3♂, Coll. Aquib Majeed, Zahid Hussain, Tabraq Ali, (in Museum, Department of Zoology, BGSB University Rajouri (J & K), India.

 Distribution.

Thailand, India.

 Note.

According to Malicky et al. (2000), D. diespiter shows similarities to D. palmipes Ito, 1986 in the morphology of segment X, although the middle cone of segment X (actually the apex of tergum IX) is significantly shorter in D. palmipes. Moreover, D. palmipes possesses bifurcate parameres. D. daidalion Malicky & Prommi, 2000 is even more similar, but it also exhibits a considerably shorter dorsal cone of tergum IX.

 Animalia Trichoptera Lepidostomatidae 463383AA-B9EA-51FE-BF7A-3522E40B932C Lepidostoma kamba (Mosely, 1939b) Dinarthrum kamba Mosely, 1939b: 338, pl 5 f 1–5. Description.

Adult ♂ (Fig. 2A–E): Male antennal scape 3.3 times as long as head, with two basal processes, proximal process long, curved distad and distal process short, digitate. Maxillary palps each two-segmented, basal segment setose and twice as long as distal segment, distal segment curved away from head. Labial palps each three-segmented, second segment twice as long as basal segment and shorter than third (2C). Forewings each covered with scales; apical forks I and II present, all forks sessile (Fig. 2E).

10.3897/contrib.entomol.73.e109883.figure2 8E8D343C-B821-5330-9343-9D0B6C5F9380

Lepidostoma kamba (Mosely, 1939b). A, B. Genitalia: A. Left lateral; B. Dorsal. C. Head with scape; D. Phallic apparatus, lateral; E. Right forewing, dorsal. BP = two basal processes of antennal scape (paired); IFA = inferior appendage (paired); IX seg = segment IX; PA = phallic paramere (paired); PH = phallus; X seg = segment X.

https://binary.pensoft.net/fig/941756

Genitalia (Fig. 2A, B, D): Segment IX in dorsal view (Fig. 2B) apically round beyond which segment X deeply excised, divided into two equal rectangles segments; segment X in lateral view (Fig. 2A) extended posteroventrad with a round apex, vertical ridge suspended below middle of segment X. Inferior appendages each two-segmented: basal segment subrectangular at base except slightly bulging in middle, small triangular forks arising from upper and inner margins of basal segment; terminal segment slightly excised at the apex. Parameres each pointed at apex, placed diagonally across genitalia; phallus cylindrical, curved, irregularly round shaped at apex, directed downwards (2D).

10.3897/contrib.entomol.73.e109883.figure3 BBBA7CFE-33A3-5040-A357-3EAD019A41C3

Collection sites for Lepidostoma kamba (Mosely, 1939b) and Lepidostoma diespiter (Malicky & Saengpradab, 2001). A. Chamoli; B. Andretta.

https://binary.pensoft.net/fig/941757 10.3897/contrib.entomol.73.e109883.figure4 22430E6D-2558-515E-B4F2-F7B32244F40E

Examples of human activities causing poor water quality and deteriorating habitat. A. Tourism; B. Urbanization; C. Mining; D. Dry streams; E. Construction of bridges.

https://binary.pensoft.net/fig/941758

Female. Not known.

 Holotype repository.

Stockholm Museum.

 Material examined.

India (new record): Uttarakhand: Chamoli, 30°26'35.8"N, 79°19'11.2"E, 13-vi-2022, 5♂, Coll. Tabraq Ali Zahid Hussain, Aquib Majeed; (in Museum, Department of Zoology, BGSB University Rajouri (J & K) India.

 Distribution.

Burma, India.

 Animalia Trichoptera Lepidostomatidae 66D013FB-3A1E-5A52-95B1-4553D92F1C5C Lepidostoma sonomax (Mosely, 1939) Dinarthrum sonomax Mosely, 1939. Holotype repository.

(Mosely 1939); Sonamarg, 28-ix-1932 (Yale North India Expedition), G. E. Hutchinson. (British Museum).

 Material examined.

India: Uttarakhand (new record); Naugoun. 30°47'26.5"N, 78°08'18.2"E 1061 m, 07-vi-2022; ♂ 5; Collectors: Zahid Hussain, Aquib Majeed, Tabraq Ali, (in Museum, Department of Zoology, BGSB University Rajouri (J & K) India.

 Distribution.

China: Tibet; India: Jammu & Kashmir, Uttarakhand.

Additional material of other <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lepidostoma">Lepidostoma</tp:taxon-name-part></tp:taxon-name></italic> species examined during the expeditions from 2019–2023

Fig. 5

 Animalia Trichoptera Lepidostomatidae 4E758CEF-338D-50F2-97B6-4CFEB1A3F47A L. inerme (McLachlan, 1878) Dinarthrum inerme McLachlan, 1878: 5–6, figs 1–4, ♂. Material examined.

India: Jammu & Kashmir: Badusgam, 33°38'24.9"N, 75°13'23.1"E, 15-viii-2-2022, 16♂, 5♀; Verinag, 33°32'10.6"N, 75°14'56.0"E, 16-viii-2-2022 4♂; Chatpal, 33°47'11.5"N, 75°23'20.7"E, 17-viii- 2022, 4♂: Himachal Pradesh: Barrot, 32°00'21.5"N, 76°51'38.8"E, 01-VI-2022, 2♂; Banjar, 31.65132, 77.33472, 02-VI-2022, 3♂, Coll. Tabraq Ali Zahid Hussain, Aquib Majeed; (in Museum, Department of Zoology, BGSB University Rajouri (J & K) India.

 Animalia Trichoptera Lepidostomatidae 978F4E3D-53D8-5557-A9CC-EB5BD992E50E L. parvulum (McLachlan, 1871) Mormonia parvula McLachlan, 1875:33, pl. 3 figs. 2–2c, ♂. Material examined.

India: Jammu & Kashmir: Achabal: 33°41'00.3"N, 75°13'19.3"E, 22-VI-2021, 12♂; Verinag: 33°32'10.6"N, 75°14'56.0"E, 22-VI-2021, 15♂; Coll. Zahid Hussain, Aquib Majeed, Tabraq Ali; (in Museum, Department of Zoology, BGSB University Rajouri (J & K) India.

10.3897/contrib.entomol.73.e109883.figure5 B78B27F8-BA9B-54BC-B281-35A96C52F272

Map: Sampling sites along with collected species during expedition in Northwest Himalayas of India.

https://binary.pensoft.net/fig/941759 Animalia Trichoptera Lepidostomatidae 97666442-F396-5F7E-B323-44C9B42700B6 L. kurseum (Mosely, 1949a) Adinarthrum kurseum Mosely, 1949: 237–238, pl. II figs. 8–12, ♂. Material examined.

India: Uttarakhand: Kampty fall: 30°29'07.5"N, 78°02'05.1"E, 07-VI-2022, 2♂; Coll. Tabraq Ali Zahid Hussain, Aquib Majeed; (in Museum, Department of Zoology, BGSB University Rajouri (J & K) India.

 Animalia Trichoptera Lepidostomatidae 991DB744-A42C-56D9-9B96-386A60A17697 L. tesarum (Mosely, 1949b) Goerodella tesarum Mosely, 1949: 421–422, pl.12 figs. 152–156, ♂. Material examined.

India: Jammu & Kashmir: Gabbar, 33°23'52.7"N, 74°38'39.8"E, 23-IX-2020, 2♂, Coll. Tabraq Ali Zahid Hussain, Aquib Majeed; (in Museum, Department of Zoology, BGSB University Rajouri (J & K) India.

 Animalia Trichoptera Lepidostomatidae AB6C96C4-ED37-53B0-A434-73E7C0A6077B L. nagana (Mosely, 1939) Dinarthrum nagana Mosely, 1939: 338–339, pl. 6 figs. 1–5, ♂. Material examined.

India: India: Jammu & Kashmir: Gabbar, 33°23'52.7"N, 74°38'39.8"E, 23-IX-2020, 2♂, Coll. Tabraq Ali Zahid Hussain, Aquib Majeed; (in Museum, Department of Zoology, BGSB University Rajouri (J & K) India.

This species has been observed/collected in India after eight decades.

 Animalia Trichoptera Lepidostomatidae 331B1FD9-FDC5-56B2-B711-E8EA27923942 L. truncatum Parey & Saini, 2012 Material examined.

India: Himachal Pradesh: Panchpulla, 32°31'41.6"N, 75°59'24.8"E, 1999 m, 28-v-2022, 2♂, Coll. Tabraq Ali Zahid Hussain, Aquib Majeed; (in Museum, Department of Zoology, BGSB University Rajouri (J & K) India.

 Animalia Trichoptera Lepidostomatidae 51B24838-24AC-548C-BBBD-7ED078C9FEB5 L. ylesomi (Weaver, 1941) Adinarthrella brunnea Mosely, 1941: 776, pl 6 f 1–5 (preoccupied in Lepidostoma by Crunoeciella brunnea Ulmer, 1905). Material examined.

India: Uttarakhand: Chamoli, 30°26'35.8"N, 79°19'11.2"E, 13-VI-2022, 2♂, Coll. Tabraq Ali, Zahid Hussain, Aquib Majeed; (in Museum, Department of Zoology, BGSB University Rajouri (J & K) India.

Checklist of Indian <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lepidostoma">Lepidostoma</tp:taxon-name-part></tp:taxon-name></italic> species

1. L. ahlae Parey & Saini, 2012

Distribution. India (Himachal Pradesh).

2. L. armatum (Ulmer, 1905)

Distribution. Nepal; India (Assam, Meghalaya).

3. L. assamense (Mosely, 1949)

Distribution. Nepal; Bhutan; India (Meghalaya).

4. L. betteni (Martynov, 1936)

Distribution. India (West Bengal, Sikkim,).

5. L. brueckmanni (Malicky & Chantaramongkol, 1994)

Distribution. Thailand; India (Assam, Meghalaya, Uttarakhand).

6. L. curvatum Parey & Saini, 2013

Distribution. India (Arunachal Pradesh).

7. L. destructum (Ulmer, 1905)

Distribution. Bhutan; India (West Bengal, Arunachal Pradesh, Assam).

8. L. digitatum (Mosely, 1949)

Distribution. India (Meghalaya).

9. L. diespiter (Malicky & Sangpradub, 2001)

Distribution. Thailand; India: Himachal Pradesh.

10. L. divaricatum (Weaver, 1989)

Distribution. Indonesia; India (Himachal Pradesh, Uttarakhand, Meghalaya, Manipur).

11. L. dirangense (Saini & Parey, 2011)

Distribution. India (Arunachal Pradesh)

12. L. doligung (Malicky, 1979)

Distribution. Indonesia; China; India (Andaman & Nicobar).

13. L. dubitans (Mosely, 1949)

Distribution. India (Meghalaya).

14. L. ferox (McLachlan,1871)

Distribution. India (Himachal Pradesh, Uttarakhand).

15. L. fuscatum (Navas,1932)

Distribution. India (Karnataka).

16. L. garhwalense Parey & Saini, 2012

Distribution. India (Uttarakhand).

17. L. heterolepidium (Martynov, 1936)

Distribution. Bhutan; Nepal; India (Uttarakhand, West Bengal).

18. L. himachalicum Saini & Parey, 2011

Distribution. India (Himachal Pradesh).

19. L. inequale (Martynov, 1936)

Distribution. Bhutan; India (Uttarakhand,Tamil Nadu).

20. L. inerme (McLachlan, 1878)

Distribution. China; India (Jammu and Kashmir, Himachal radesh).

21. L. kamba (Mosely, 1939b)

Distribution. Burma; India: Uttarakhand.

22. L. kashmiricum Saini & Parey, 2011

Distribution. India (Jammu & Kashmir, Sikkim, West Bengal).

23. L. kjeri Parey & Pandher, 2019

Distribution. India (Uttarakhand, Arunachal Pradesh).

24. L. khasianum (Mosely, 1949c)

Distribution. India (Meghalaya, Tamil Nadu).

25. L. kimsa (Mosely, 1941)

Distribution. India (Sikkim).

26. L. kurseum (Mosely, 1949)

Distribution. Nepal; India (Sikkim, Meghalaya, Himachal Pradesh).

27. L. lanca (Mosely, 1949)

Distribution. India (Karnataka).

28. L. latum (Martynov, 1936)

Distribution. India (Jammu & Kashmir, Himachal Pradesh, West Bengal.

29. L. libitana (Malicky, 2003)

Distribution. Bhutan; India (Himachal Pradesh).

30. L. liber (Malicky, 2007)

Distribution. Bhutan; India (Arunachal Pradesh).

31. L. lidderwatense Parey, Morse & Pandher, 2016

Distribution. India (Jammu & Kashmir).

32. L. margula (Mosely, 1949)

Distribution. India (Jammu & Kashmir).

33. L. mechokaense Parey & Saini, 2013

Distribution. India (Arunachal Pradesh).

34. L. moulmina (Mosely, 1949)

Distribution. India (Assam, Meghalaya)

35. L. nagana (Mosely, 1939)

Distribution. India (Jammu & Kashmir, Himachal Pradesh).

36. L. nubragangai Dinakaran, 2013

Distribution. India (Tamil Nadu)

37. L. palmipes (Ito, 1986)

Distribution. Nepal; China; India (Uttarakhand, Arunachal Pradesh, Sikkim).

38. L. palnia (Mosely, 1949)

Distribution. India (Tamil Nadu)

39. L. parvulum (McLachlan, 1871)

Distribution. Uzbekistan; India (Jammu & Kashmir).

40. L. punjabicum (Martynov, 1936)

Distribution. India: (Himachal Pradesh,Uttarakhand)

41. L. sainii Parey, Morse & Pandher, 2016

Distribution. India (Uttarakhand, Himachal Pradesh, Meghalaya).

42. L. serratum (Mosely, 1949c)

Distribution. United States of America (USA); India (Meghalaya, Assam).

43. L. sika (Mosely, 1949)

Distribution. India (Arunachal Pradesh, Sikkim).

44. L. simplex (Kimmin, 1964)

Distribution. Nepal; India (Uttarakhand).

45. L. sonomax (Mosely, 1939)

Distribution. Tibet; India (Jammu & Kashmir).

46. L. sonmargae Parey & Saini, 2012

Distribution. India (Jammu & Kashmir).

47. L. steelae (Mosely, 1941)

Distribution. India (Meghalaya).

48. L. tesarum (Mosely, 1949)

Distribution. Bhutan (Malicky 2007); India (Himachal Pradesh, Uttarakhand).

49. L. trilobatum Parey, Morse & Pandher, 2016

Distribution. India (Arunachal Pradesh).

50. L. truncatum Parey & Saini, 2012

Distribution. India (Himachal Pradesh).

51. L. ylesomi (Weaver, 1941)

Distribution. Nepal; India (Sikkim, Uttarakhand, Jammu & Kashmir).

 Discussion

During the survey, (Fig. 5) we documented only ten species of Lepidostomatidae, all belonging to the genus Lepidostoma. With the addition of two new records, L. diespiter (Malicky & Saengpradab, 2001) and L. kamba (Mosely, 1939b), from Himachal Pradesh and Uttarakhand respectively in the L. ferox Branch, the number of Indian species of Lepidostoma is now 51 and the number of Indian Lepidostomatidae species is 65. Moreover, during our expedition L. sonomax was collected for the first time from Uttarakhand, which was reported previously only from Kashmir by Mosely in early 1939.

 Conclusion

The streams and water bodies from where the other species specimens were investigated and documented previously (Saini et al. 2011; Parey et al. 2012, 2013, 2016, 2019; Dinakaran et al. 2013) were either polluted or dry due to increasing global temperature or shifting of precipitation. Poor water quality and deteriorating habitats directly limit the presence of Lepidostomatidae, other caddisflies, and other freshwater biota (e.g., Houghton and DeWalt 2023). For example, urbanization, the use of chemicals in agricultural fields, construction of bridges and dams persistently threaten freshwater species (Dudgeon 2019). The construction of dams causes the fragmentation of water bodies that disrupts the species’ habitat (Madadi et al. 2017). Pesticides used in agricultural fields contaminate soil and water and disrupt the species’ existence by flowing into water bodies (Rajmohan et al. 2020). Recreational bathing by tourists in many freshwater bodies is another major driver that acts as a catalyst in pollution and contamination of these water reservoirs by discharging litter and sewage that negatively affects the habitat of the Lepidostomatidae as well as other aquatic organisms (Bhatnagar et al. 2016). Adverse effects on Lepidostomatidae are also attributed to mining which alters the course of water and pollutes it (Liu Y. et al. 2022) (Figs 4A–E).

 Acknowledgments

We thank Dr. Karl Kjer for the ultraviolet light used in this research. The corresponding author is thankful to the Science and Engineering Research Board (SERB), Government of India for providing a financial grant vide sanction order no. CRG/2020/001494 Dated 1-12-2020 and providing travel grant under ITS (International Travel Support) scheme with file no. ITS/2022/001567 dated 27 July 2022, to attend the XVII International Symposia on Trichoptera at Lunz am See, Austria. Thanks to Dr. Simon Vitecek for providing best possible facilities to the corresponding author at Lunz Amm See, Austria. Thank you, Dr. John C. Morse, for the comments that improved this manuscript. We also thank anonymous reviewers for critically reading this manuscript.

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