Journal of Threatened Taxa | | 26 March 2021 | 13(3): 18045–18049


ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print)

#6517 | Received 04 August 2020 | Final received 12 October 2020 | Finally accepted 06 March 2021



Habitat association and hybridization in woodbrowns (Lethe nicetas, L. sidonis, & L. dakwania) (Lepidoptera: Nymphalidae: Satyrinae) in Kedarnath Musk Deer Reserve, western Himalaya


Arun Pratap Singh 1 & Tribhuwan Singh 2


1,2 Entomology Branch, Forest Protection Division, Forest Research Institute (ICFRE), P.O. New Forest, Dehradun, Uttarakhand248006, India.

 1 (corresponding author), 2




Editor: Anonymity requested.   Date of publication: 26 March 2021 (online & print)


Citation: Singh, A.P. & T. Singh (2021). Habitat association and hybridization in woodbrowns (Lethe nicetas, L. sidonis, & L. dakwania) (Lepidoptera: Nymphalidae: Satyrinae) in Kedarnath Musk Deer Reserve, western Himalaya. Journal of Threatened Taxa 13(3): 18045–18049.


Copyright: © Singh & Singh 2021. Creative Commons Attribution 4.0 International License.  JoTT allows unrestricted use, reproduction, and distribution of this article in any medium by providing adequate credit to the author(s) and the source of publication.


Funding: Indian Council of Forestry Research & Education (ICFRE).


Competing interests: The authors declare no competing interests.


Acknowledgements: The current findings are part of a ICFRE research project entitled “Butterflies associated with different forest types/sub-types in Uttarakhand (FRI-627/FED-44; 2017-2021)”  being  carried out at the Forest Research Institute, Dehradun.



The ‘woodbrown’ group of the genus Lethe (Nymphalidae: Satyrinae: Satyrini: Lethina) occurs as five species in western Himalaya.  The most common and widely occurring species is the Common Woodbrown Lethe sidonis (Hewitson, 1863) (45–60 mm) which occurs from Chamba, Himachal Pradesh up to Arunachal Pradesh and Shan states in Myanmar from April to October at 975–3,352 m.  Its larva is known to feed on Hill Bamboo Arundinaria falcata Nees.  A similar looking and lesser known species is the White-wedged Woodbrown or the Garhwal Woodbrown Lethe dakwania Tytler, 1939 found in Garhwal.  Specimens of both the sexes of this little known species were collected by H.C. Tytler during August 1914 from Dakwani, eastern Garhwal (2,700m) in northern India.  Another species that occurs along with these two is the Yellow Woodbrown Lethe nicetas (Hewitson, 1863) (48–55 mm) which is distributed from Kangra in Himachal Pradesh up to Arunachal Pradesh in the Himalaya, northeastern India and northeastern part of Myanmar.  It occurs at 1,700–2,620 m with a flight period from May to November and is ‘not rare’ in its distribution range.  This species is more common in June–October (900–1,800 m) in Kumaon region of the western Himalaya.  The fourth species is the Himalayan Barred Woodbrown Lethe maitrya maitrya de Nicéville, [1881] (45–55 mm) which occurs from Kullu in Himachal Pradesh up to Sikkim and Bhutan where it is ‘not rare’ at 2,500–3,800 m in April–October.  The fifth species is the Scarce Woodbrown Lethe siderea siderea Marshall, 1881 (48–55mm) that is distributed from Garhwal to northeastern India & northern Burma where it is ‘rare’ and occurs between 2,000–2,620 m from May–October (Mackinnon & Nicéville 1899; Hannyngton 1910; Evans 1932; Wynter-Blyth 1957; Smith 1989, 2006; Varshney & Smetacek 2015; Singh & Sondhi 2016; Kehimkar 2016; Gasse 2013).

During the course of several surveys carried out in Kedarnath Musk Deer Reserve (KMDR) in 2006–2019, observations were recorded and random samples collected of the Lethe genus of the group ‘Woodbrown’ at various locations representing different altitudes and vegetation types.  Analysis of photographs, specimens and male genitalia revealed the occurrence of only three species of woodbrowns in KMDR out of five known from western Himalaya.  These were: L. sidonis, L. dakwania, & L. nicetas (Image 1, 2, 3, 5, 6 & 7).  The species have been earlier reported from Mandal and Kanchula Kharak areas inside KMDR as “common” (Singh & Sondhi 2016).  Upon examination, these species showed distinct male genitalia.  In L. sidonis, the uncus, as seen from the side is distinctly raised, and higher and thicker in the middle, and then sharply bent downwards (Image 1 a–c).  In L. dakwania the uncus is evenly curved and not thicker and raised in the middle, or suddenly bent downwards (Image 2 a–c) (Tytler 1939).  In L. nicetas (Image 3 a–c) the uncus is bent sharply downwards in the beginning without being thicker or raised in the middle.  While in L. maitrya (specimen collected from Mussoorie, Garhwal) the uncus is not bent at all but straight and held horizontally in front (Image 4 a–b).  Two specimens collected from KMDR seemed morphologically quite similar to L. nicetas but were distinct as they had yellow markings on the under and upper forewing and under hindwing being ‘more extensive’ than in L. nicetas (Image 8 & circled portions of upper forewings- Image 3 & 9).  These specimens were dissected for examining their genitalia but did not reveal any genital organ and were thus classified as hybrids.  These specimens were collected during 2006 and then again during 2017, which suggests that the phenomenon of hybridization in an ongoing process in this part of KMDR.

Examination of the altitudinal distributional and forest type association (Figure 1 & Table 1) of these three species in the study area revealed that L. nicetas is associated with 12/C1b Moru oak forest (Champion & Seth 1968) and mainly occurs in abundance at 2,260–2,402 m.  On the other hand L. dakwania occurred at a much higher elevation at 2,729–2,765 m and showed association with mainly 12/C1d western mixed coniferous forest.  While L. sidonis had a much wider altitudinal distribution range at 1,700–2,600 m and occurred in at least three forest types: 12/C1a Ban Oak forest, 12/C1b Moru Oak forest, and also 12/C1d western mixed coniferous forest, thus sharing common forest-type habitat with both nicetas and dakwania in KMDR, therefore having greater chances of hybridization with L. nicetas.  The hybrids collected (Image 8 & 9) are most likely to be between nicetas and sidonis.  The current findings call for more research into the matter.



Table 1. Plant species* (trees & bamboos) composition of the three different forest sub-types (Champion & Seth 1968--) associated with the three species of Woodbrowns (Lethe nicetas, L. sidonis, & L. dakhwania) in Kedarnath Musk Deer Reserve, western Himalaya.



12/C1a Ban Oak forest

(Quercus leucotrichophora A.Camus)

12/C1b Moru Oak forest

(Quercus floribunda Lindl. ex A.Camus)

12/C1d western mixed coniferous forest

(Abies pindrow (Royle ex D.DonRoyle,

Pinus wallichiana A.B.Jacks)

a) Trees


b) Dwarf Bamboos

a) Acer caesium Wall. ex Brandis 

Acer oblongum Wall. ex DC. 

Aesculus indica (Wall. ex Cambess.) Hook.

Alnus nepalensis D.Don

Betula alnoides Buch.-Ham. ex D.Don

Boehmeria rugulosa Wedd.

Cinnamomum tamala (Buch.Ham.) T.Nees & C.H.Eberm

Cornus capitata Wall. ex Roxb

Euonymus lacerus Buch.-Ham.

Ficus auriculata Lour.

Fraxinus micrantha Lingelsh.

Inula cuspidate (Wall. ex DC.) C.B.Clarke

Lindera pulcherrima (Nees) Benth.

Litsea umbrosa (Nees) Nees

Lyonia ovalifolia (Wall.) Drude,

Machilus odratissima Nees

Machilus duthiei King ex J.D.Hooker

Marsine semiserata Wallich

Pryrus pashia Linnaeus

Rhododendron arboretum Sm.

Sarcococca saligna (D.Don) Müll.Arg.

Xanthoxylum armatum DC.


b) Sinarundinaria falcata (Nees) C.S.Chao & Renvoize

a).  Acer caesium Wall. ex Brandis 

Acer sterculiaceum Wall. 

Aesculus indica (Wall. ex Cambess.) Hook.

Betula alnoides Buch.-Ham. ex D.Don

Carpinus viminea Wall. ex Lindl.

Eurya acuminata DC.

Fraxinus micrantha Lingelsh.

Ilex dipyrena Wall.

Machilus duthiei King ex J.D.Hooker

Rhamnus purpureus Edgew.

Rhododendron arboreum Sm.

Symplocos chinensis (Lour.) Druce




b).  Sinarundinaria falcata (Nees) C.S.Chao & Renvoize

a).   Quercus semecarpifolia Sm.

Acer caesium Wall. ex Brandis 

Acer cappadocicum Gled

Euonymus lacerus Buch.-Ham

Rhdodendron aboreum Sm.

Rhamnus purpureus Edgew.

Smilax vaginata Decne.

Taxus wallichiana Zucc.

Juniperus indica Bertol.


b). Thalmnocalamus falconeri Hook.f. ex Munro

Yushania anceps (Mitford) W.C.Lin


Identification of plant species based in the field with the help of field guide (Rai et al. 2017) and herbarium specimens collected during field

surveys by the authors and identified at FRI, Dehradun Herbarium with the help of plant taxonomists.


For figure & images - - click here





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