Journal of Threatened
Taxa | www.threatenedtaxa.org | 26 September 2024 | 16(9): 25843–25855
ISSN 0974-7907
(Online) | ISSN 0974-7893 (Print)
https://doi.org/10.11609/jott.8485.16.9.25843-25855
#8485 | Received 19
April 2023 | Final received 22 July 2024 | Finally accepted 10 September 2024
Taxonomic review of genus Gazalina
Walker (Thaumetopoeinae: Notodontidae: Lepidoptera) from India
Amritpal Singh Kaleka 1 ,
Gagan Preet Kour Bali 2 &
Navkiran Kaur 3
1,3 Department of Zoology &
Environmental Sciences, Punjabi University, Patiala, Punjab 147002, India.
2 Department of Biosciences-UIBT,
Chandigarh University, Mohali, Punjab 140413, India.
1 apskaleka@gmail.com, 2 gaganviren@gmail.com
(corresponding author), 3 navkiran.dandiwal@gmail.com
Editor: Jatishwor Singh Irungbam, Centrum ALGATECH,
Třeboň, Česká Republika. Date of publication: 26 September
2024 (online & print)
Citation: Kaleka, A.S., G.P.K. Bali & N. Kaur (2024). Taxonomic
review of genus Gazalina Walker (Thaumetopoeinae: Notodontidae:
Lepidoptera) from India. Journal of Threatened Taxa 16(9): 25843–25855. https://doi.org/10.11609/jott.8485.16.9.25843-25855
Copyright: © Kaleka et al. 2024. 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: University Grants Commission, New Delhi (MANF Scheme);
F1-17.1/2O13-14/MANF/2013-14-SIK-27073 Dated: 6-02-2014.
Competing interests: The authors declare no competing interests.
Author details: Amritpal
Singh Kaleka is presently serving in the Department of Zoology & Environmental Sciences, Punjabi University
and has 31 years of teaching and research experience. He successfully completed
research projects from CSIR, DST, UGC and currently a research project from
MoEFCC is in progress. He published 114 research papers, 50
articles and book chapters. He attended 60 Conferences and remained
instrumental in organizing 16 conferences/Symposia and workshops. Dr.
Gagan Preet Kour Bali is currently affiliated with Chandigarh
University, Mohali, Punjab as an assistant professor. Her area
of specialization is entomology and she is working to assess moth diversity
from north-west Himalayas since 2014. She has contributed 16 research papers in
scientific Journals of repute. She is also associated with Intech-Open as an author & reviewer and contributed five book chapters
pertaining to insects. Navkiran
Kaur affiliated with Punjabi University, Patiala. Her area of interest is
entomology and has worked on internal genitalic features of moths.
Author contributions: Study design and article
drafting—ASK & GPKB; molecular analysis—GPKB; Lab & field work—ASK,
GPKB & NK.
Acknowledgements: The authors are thankful to the
authorities of forest departments of Himachal Pradesh, and Jammu & Kashmir
for their support during field surveys for sample collection.
Abstract: The taxonomic position of the
genus Gazalina Walker remained ambiguous and it continuously reshuffled
its position in the family Notodontidae and subfamily Lymantriinae of family
Erebidae. The present study has been carried out to elucidate its taxonomic
position in accordance with the morphological, molecular, larval, and
behavioural characteristics of its species. A detailed account of two species
namely Gazalina chrysolopha (Kollar) and Gazalina apsara (Moore),
the type species of this genus, has also been given. The CoI sequence of Gazalina
chrysolopha (MH795522) and Gazalina apsara (MH790288) from the
present study were compared with the available reference sequences in Genbank
by using BLAST. Its placement under the subfamily Thaumetopoeinae of the family
Notodontidae has been justified.
Keywords: Apsara, Chrysolopha,
diagnosis, genitalia, Lymantriidae, phylogenetic, processionary moths, sequence
analysis, taxonomic, wing venation.
Introduction
Walker (1865) established the
genus Gazalina Walker 1865 with Gazalina venosata Walker
as its type species from northern India
under the family Liparidae. Kirby (1892) followed the same nomenclature.
Hampson (1892) reported three species namely G. apsara (Moore, 1859), G.
chrysolopha (Kollar, 1844) and transversa Moore, 1879 from India.
While dealing with Eastern and Australian moths, Swinhoe (1900) described a new
species, Gazalina intermixta, under this family. Grunberg (1912) shifted
this genus to the family Notodontidae. Later, Swinhoe (1922) revised various
genera referable to the family Liparidae and discussed four species—G.
apsara Moore (formerly Venosata Walker), G. chrysolopha
(Kollar), G. intermixta Swinhoe, and G. transversa Moore—under
the family Liparidae. Kiriakoff (1968) further placed Gazalina in the
family Thaumetopoeidae, a distinct family.
Further, he studied the male genitalia of apsara Moore, the type
species of this genus, and diagnosed the absence of gnathos, a diagnostic
feature of the family Notodontidae (Kiriakoff 1970). Cai (1979) treated it
under the family Notodontidae and discussed three species G. apsara
Moore, G. chrysolopha (Kollar), and G. transversa Moore in
‘Economic Insect Fauna of China’. Miller (1991) again doubted the placement of
the genus Gazalina Walker. Sugi (1994) described three species—G.
apsara (Moore), G. chrysolopha (Kollar), G. transversa
Moore—from Nepal under the family Notodontidae. Raman (1998) studied the
outbreak of G. chrsolopha (Kollar) accounting for it under family
Notodontidae. Wu (2002) described four species—G. apsara (Moore), G.
chrysolopha (Kollar), G. transversa Moore, and G. putrificata Sugi—from
China under Notodontidae. Srivastava & Mukhopadhyay (2006) studied the life
cycle and bio-ecology of G. chrsolopha (Kollar) accounting for it under
the family Notodontidae. Sanyal et al. (2011) studied Gazalina apsara
(Moore) as an indicator species from the Himalaya under the family
Notodontidae. Kocak & Kemal (2016) enlisted it under the family
Thaumetopoeidae. While dealing with molecular phylogeny, Kobayashi & Nonaka
(2016) also discussed the genus Gazalina Walker in the subfamily
Thaeumatopoeinae. Uniyal et al. (2016) catalogued the genus Gazalina
Walker under the subfamily Thaumetopoeinae of the family Notodontidae from Gangotri
landscape, Uttarakhand, India. Shah et
al. (2017) enlisted the genus Gazalina Walker under the subfamily
Lymantriinae. While studying the medical complications caused by different
species of Gazalina Walker, Manandhar et al. (2018) discussed it
under the family Notodontidae. Bhattacharyya et al. (2019) enlisted the genus Gazalina
Walker under the subfamily Thaumetopoeinae of the family Notodontidae from
Neora Valley, West Bengal, India. Recently, Chettri et al. (2021) enlisted Gazalina
chrysolopha Kollar and G. transversa Moore from Sikkim and placed
these species under the family Notodontidae. Gurung et al. (2021) described Gazalina
chrysolopha Kollar as a major pest consuming foliage of trees especially Alnus
nepalensis D.Don, Rhododendron arborium Smith, and other fodder
plants; but the taxonomic position was not clear. Khanal & Shrestha (2022)
studied the diversity, distribution, and medical significance of Gazalina
species from Nepal. The morphological characters including external as well as
internal genitalic features, behavioural characters, and molecular analysis of
two species namely Gazalina chrysolopha (Kollar) and Gazalina apsara
(Moore), the type species of this genus have been compiled in detail to
elucidate the position of genus Gazalina Walker.
Material
and Methods
The adult moths were collected
from different localities of Himachal Pradesh and Jammu & Kashmir (India)
by using light traps equipped with a 160w mercury bulb and vertical white
screen and their behaviour was observed during the collection period. The
external morphological characters were studied as such from the preserved
specimens. The permanent slides were prepared to study the wing venation. The
male and female moths were dissected to examine the external and internal genitalic
features and the terminology for naming various genitalic parts given by Klots
(1970) was followed. The DNA was extracted from the preserved moth samples
using the phenol-chloroform-isoamyl alcohol method given by Sambrook et al.
(1989). The mitochondrial cytochrome oxidase subunit I (COI) gene was amplified
using the universal primer pair under standard PCR conditions (Folmer et al.
1994).
Forward –
(LCO1490:
5’-GGTCAACAAATCATAAAGATATTGG-3’)
Reverse –
(HC02198:
5’-TAAACTTCAGGGTGACCAAAAAATCA-3’)
The purified PCR products were
sequenced at Amnion Biosciences Pvt. Ltd, Sequencing Dept, #112, Doddenna industrial
area, 16A Cross, Vishwaneedam post (D), Bangalore, Karnataka 560091, India.
The sequences were submitted to
Genbank for accession numbers as Gazalina chrysolopha MH795522 & Gazalina
apsara MH790288. Multiple sequence alignment was performed with CLUSTAL x
software and a phylogenetic tree was constructed using the maximum likelihood
method (ML) in MEGA (version 6) software. The confidence level of each branch
was evaluated through bootstrap analysis with 1000 replicates (Tamura et al.
2021).
Systematic Account
Genus Gazalina Walker 1865
Walker, 1865, List Spec.
Lepid. Insects Colln. Brit. Mus., 32: 298; Swinhoe, 1922, Ann. Mag.
Nat. Hist., 9(9)10 (58): 472.
Oligoclona Felder, 1874, Reis. Freg.
Nov., 2: pl. 94.
Ansonia Kiriakoff, 1967, in Wytsman,
Genera Insect., 217(B): 57.
Type Species:
Gazalina
apsara (Moore) = venosata
Walker
Distribution:
India; China.
Diagnosis: Medium-sized moths, usually white
in colouration. Labial palpi extremely minute. Antennae bipectinate in males,
serrate in females, and pectinations are reduced at the distal end. Forewing
with ground colour white, without any distinct markings; discal cell more than
half the length of the wing, closed; 1A+2A basally forked; 3A absent; Cu1
and M3 from lower angle of cell; M2 from lower angle of
cell; M1 just above the middle of discocellulars; R5 from
the upper angle of cell; R4-R2 stalked from the upper
angle of cell; R1 well before upper angle of cell; Sc from the base
of wing not reaching the apex. Hindwing without any distinct markings; discal
cell more than half the length of wing, closed; Cu1 and M3
from near lower angle of cell; M2 from near middle of
discocellulars; M1 and Rs stalked from upper angle of cell. Legs
dressed with white scales; fore-tibia with an epiphysis, mid-tibia with one
pair of tibial spurs; hind-tibia with two pairs of tibial spurs. Abdomen
slender, banded with black and white scales; distal segments fringed with long
white scales in males, distinct golden anal tuft in females. Male genitalia
with uncus of moderate size, gnathos represented by conjoined, triangular processes;
saccus absent; juxta well developed; valva simple, basal half broad, distal
half narrow; aedeagus of moderate size, vesica without any distinct cornuti.
Female with corpus bursae globular; signum absent; ductus bursae narrow,
medially constricted; apophysis of moderate length, both pairs with equal
length with dilated tips; papilla analis prominent, setosed; pseudo-papillae
indistinct; sterigmatic plate well developed.
Gazalina chrysolopha (Kollar, 1844)
Liparis
chrysolopha Kollar, 1844,
Hügel Kaschmir und Das Reich der Siek, 4: 470.
Gazalina
chrysolopha Kollar:
Hampson, 1892, MothsIndia, 1: 469; Swinhoe, 1922, Ann. Mag. Nat.
Hist., (9) 10 (58): 472; Wu, 1999, Fauna Sinica, 31: 832.
Dasychira
antica Walker,
1855, List Spec. Lepid. Insects Colln. Brit. Mus., 4: 867; Swinhoe,
1922, Ann. Mag. Nat. Hist., (9) 10 (58):472.
Oligoclona
chordigera Felder, 1874,
Reis. Freg. Nov. 2 (4): 94.
Type
Locality: Kashmir,
India
Wing Expanse:
Male: 40–46
mm; Female: 52–60 mm.
Body Length: Male: 14–19 mm; Female: 18–21
mm.
Diagnosis: Forewing whitish, veins distinct
with black scales beyond the medial oblique line; vein M1 not
stalked with radial veins. Male genitalia with uncus notched distally; juxta
with distal end notched; distal end of valva rounded and produced. Internal
male genitalic organs with testis rounded; seminal vesicle-I originating from
testis separately; seminal vesicle-II sickle-shaped; ductus ejaculatorius
duplex curled; accessory glands free distally; primary simplex divided into
three sections. Internal female genitalic organs with eggs rounded, covered by
a thin translucent sheath; accessory gland ducts long.
Description: Head with vertex and frons
clothed with white scales. Labial palpi fringed with black scales. Antennae
with scape covered with white scales, flagellum black. Thorax, collar, and
tegula dressed with yellowish-white scales; underside white. Legs dressed with
white scales. Abdomen slender, furnished with black scales with white bands;
distal segments fringed with long white scales; distinct golden anal tuft in
females. Hindwing white. Forewing (Image 1) with Cu2 beyond
two-thirds of cell; Cu1 from well before lower angle of cell; M3
from lower angle of cell; M2 from middle of discocellulars; M1
from upper angle of cell; R5-R3 well stalked from upper
angle of cell; R2 absent; R1 beyond three-fourth of cell;
Sc from base of wing, not reaching to apex. Hindwing (Image 2) with Cu2
from well beyond two-thirds of the cell; Cu1 from well before the
lower angle of the cell; M3 from the lower angle of the cell; M2
from well above the middle of discocellulars; M1 and Rs well
stalked from upper angle of cell; Sc+R1 from base of wing
anastomosing at one-third of cell reaching till apex of wing.
Male
genitalia (Image 3–7): Uncus of moderate size, basal half broad, distal end notched giving
bifid appearance, setosed and more sclerotized; gnathos represented by
well-sclerotized, triangular, setosed paired projections; tegumen U-shaped,
moderately sclerotized, both arms of equal width, longer than vinculum;
vinculum V-shaped, weakly sclerotized, without any distinct saccus; juxta
moderately sclerotized, medially dilated on lateral sides, distal end notched.
Valva simple, broad, rounded; moderately sclerotized; setosed; distal end
produced on costal side with round, setosed tip. Aedeagus long, narrow;
proximal half flap-like; ductus ejaculatorius entering under this flap; distal
half tubular; distal end produced with an extension having serrations; walls of
aedeagus near distal end with minute denticles; vesica without any distinct
armature. The internal male genitalic (Image 1) attributes along with the
morphometric analysis have been studied in detail (Table 1).
Female
genitalia (Image 8–9): Corpus bursae large, circular, membranous; signum absent; ductus bursae
membranous, narrow, entering into well sclerotized oblong antrum; ductus
seminalis originating from the middle of ductus bursae; apophysis of moderate
length, semi-sclerotized, posterior apophysis slightly longer than anterior
apophysis, both pairs with prominently dilated apices; papilla analis ovoid,
well setosed with short and long setae; sterigmatic plate rectangular, notched
medially, well sclerotized. The internal genitalic (Image 2) details are
tabulated in Table 2.
Material
examined: India:
Himachal Pradesh: Baijnath, 998m, 32.052°N & 76.648°E, 09.x.2013, 1♂; Baila, 1,520 m, 31.056°N &
76.831°E, 04.vi.2014, 1♂; Basantpur, 2,148 m, 31.208°N & 77.174°E, 09.vii.2013, 1♂; 10.vii.2013, 7♂♂, 8♀♀; Dalhousie, 1,970 m, 32.587°N & 75.971°E, 07.vi.2013, 1♂, 2♀♀; Dilman, 1,552 m, 30.824°N &
77.134°E, 11.viii.2013, 12♀♀; 03.ix.2015, 4♂♂, 2♀♀; Fagu, 2,533 m, 31.085°N &
77.300°E, 04.ix.2016, 4♂♂, 3♀♀; Habban, 2,063 m, 30.915°N & 77.325°E, 07.vii.2014, 1♀; 07.ix.2015, 4♂♂, 4♀♀; Hadsar, 2,300 m, 32. 455°N & 76.613°E , 11.vi.2013, 1♂; Khajjiar, 1,920 m, 32.555°N & 76.065°E, 08.vi.2013, 1♂; Kharouth, 1,300 m, 32.065°N & 76.450°E, 09.vi.2017, 5♂♂, 3♀♀; Naina Tikkar, 1,552 m, 30.804°N
& 77.119°E, 01.09.2015, 4♂♂, 4♀♀; Narkanda, 2,708 m, 31.257°N & 77.460°E, 17.vi.2013, 1♂; Nauni, 1,275 m, 30.860°N & 77.173°E, 07.viii.2013, 1♀; Sabathu, 1,265 m, 30.975°N & 76.990°E, 09.viii.2013, 1♂; 15.vii.2016, 4♂♂, 2♀♀; Serighat, 1,520 m, 31.050°N & 77.069°E, 10.viii.2013, 54♂♂, 10♀♀; 26.vi.2017, 5♂♂, 5♀♀; Theog, 1,965 m, 31.118°N &
77.359°E, 20.vi.2014, 1♂, 1♀; 08.ix.2016, 4♂♂, 3♀♀; Jammu & Kashmir: Batote, 1,560 m, 33.121°N &
75.32°E,11.vii.2014, 1♀; Hote, 400 m, 32.825°N &
75.641°E, 04.ix.2013, 2♀♀, Lamberi, 336 m, 33.130°N &
74.260°E, 11.ix.2013, 1♂, 2♀♀. Coll.: Gagan Bali & Navkiran Kaur.
Distribution:
India:
Himachal Pradesh, Jammu & Kashmir, Sikkim, West Bengal; China.
Comments: Kollar (1844) originally
described this species under the genus Liparis Ochsenheimer from
Kashmir. Hampson (1892) transferred it to the present genus. Haruta (1993)
collected crysolopha Kollar from Godawari (1,600 m) southeastern
Kathmandu. During another expedition, the species was recorded from Dagchu
(2,880 m) and Jin (2,340 m) in eastern Nepal (Haruta, 1994). Shah et al. (2017)
reported this species from West Bengal. Chettri et al. (2021) enlisted Gazalina
chrysolopha Kollar from the Tadong region of Sikkim under the family
Notodontidae. Dewan et al. (2022) recorded chrysolopha Kollar in the
Trans Himalayan region of western Nepal and placed it under the family
Notodontidae.
Gazalina apsara (Moore, 1859)
Dasychira
apsara Moore, 1859,
Cat. Lepid. Ins. Mus. Nat., 2: 341.
Gazalina
apsara Moore:
Hampson, 1892, Moths India, 1: 468-469; Swinhoe, 1922, Ann. Mag. Nat.
Hist., (9) 10 (58): 472; Wu, 1999, Fauna Sinica, 31: 830–832.
Gazalina
venosata Walker,
1865, List. Spec. Lepid. Insects Colln. Brit. Mus., 32: 398; Swinhoe,
1922, Ann. Mag. Nat. Hist. (9) 10 (58): 472.
Oligoclona
nervosa Felder and
Rogenhofer, 1875; Reis. Freg. Nov., 2(4): 95.
Type
Locality: Northern
India.
Wing Expanse: Male: 40–42 mm; Female: 54 mm.
Body Length: Male: 14–17 mm; Female: 19–21 mm.
Diagnosis: Forewing whitish, veins without
black scales; vein M1 stalked with radial veins from the upper angle of the
cell. Male genitalia with uncus having a curved hook-like distal end; juxta
with distal end curved; distal end of valva beaked. Internal male genitalic
organs with testis ellipsoidal; seminal vesicle-I originating from the testis
in the fused state; seminal vesicle-II spindle-shaped; ductus ejaculatorius
duplex comma-shaped; accessory glands fused distally; primary simplex divided
into four sections. Internal female genitalic organs with eggs are rectangular,
without any covering; accessory gland ducts are small.
Description: Head with vertex and frons
clothed with white scales. Labial palpi fringed with black scales. Antennae
with scape covered with white scales, flagellum black. Thorax, collar, and
tegula dressed with yellowish-white scales; underside white. Legs dressed with
white scales. Abdomen slender, furnished with black scales with white bands;
distal segments fringed with long white scales; distinct golden anal tuft in
females. Hindwing white. Forewing (Image 10) with Cu2 from beyond
two-thirds of cell; Cu1 well before lower angle of cell; M3
from lower angle of cell; M2 just above middle of discocellulars; M1,
R5-R3 stalked from upper angle of cell; R2
absent; R1 well before upper angle of cell; Sc from base of wing,
not reaching apex. Hindwing (Image 11) with Cu2 from well before
two-thirds of the lower angle of the cell; Cu1 from well before the
lower angle of the cell; M3 from lower angle of cell; M2
from well above middle of discocellulars; M1 and Rs well stalked
from upper angle of cell; Sc+R1 from base of wing anastomosing with
cell well before its middle.
Male
genitalia (Image 12–16): Uncus small, basal half oval, distal half narrow, curved, hook-like,
dorsally setosed with short setae, well sclerotized; gnathos represented by
small, conjoined triangular processes, setosed with short setae; tegumen
moderately sclerotized, both arms dilated laterally beyond middle towards
vinculum; vinculum moderately sclerotized, narrow, without distinct saccus;
juxta large, oblong, dome-shaped, well sclerotized, distal end having a well
sclerotized nearly squarish, curved projection. Valva simple, well sclerotized,
basal half broad, saccular margin produced, setosed, distal half narrow, distal
end produced on dorsal margin giving weakly bifid appearance. Aedeagus of
moderate length, well sclerotized; proximal half flap-like; distal end bifid,
one wedge-shaped and other with prominent serrations on inner margins; vesica
without any distinct cornuti. The internal male genitalic (Figure 3) attributes
along with the morphometric analysis have been summarized in Table 3.
Female
genitalia (Image 17–18): Corpus bursae globular, membranous; signum absent; ductus bursae
membranous, narrow, medially constricted; ductus seminalis originating from the
middle of ductus bursae; antrum well sclerotized, nearly squarish; apophysis of
moderate length, almost of equal length, prominently dilated at distal end;
papilla analis prominent, broad, well setosed with long and short setae;
pseudo-papillae not distinct; sterigmatic plate large, prominently developed.
The internal genitalic (Figure 4) details are given in the tabulated form
(Table 4).
Material
examined: India:
Himachal Pradesh: Basantpur, 2,148 m, 31.208°N & 77.174°E, 10.vii.2013, 1♂; Cheog, 2,086 m, 31.068°N & 77.312°E, 05.ix.2016, 4♂♂, 3♀♀; Fagu, 2,533 m, 31.085°N &
77.300°E, 03.ix.2016, 3♂♂, 4♀♀; Ghoond, 2,086 m, 31.075°N & 77.425°E, 01.ix.2016, 4♂♂, 3♀♀; Jhumar, 2,133 m, 32.548°N &
76.146°E,11.vii.2015, 6♂♂, 3♀♀; Mahasu, 2,086 m, 31.100°N & 77.504°E, 30.viii.2016; 3♂♂, 2♀♀; Narkanda, 1,265 m, 31.257°N
& 77.460°E, 17.vi.2013, 1♀; Serighat, 1,520 m, 31.050°N
& 77.069°E, 11.viii.2013, 1♂, 4♀♀; 25.vi.2017, 4♂♂, 3♀♀; Jammu & Kashmir: Lamberi, 336 m, 33.130°N & 74.260°E,
11.ix.2017, 1♀. Coll.: Gagan Bali &
Navkiran Kaur.
Distribution: India: Himachal Pradesh, Jammu
& Kashmir, Sikkim; China.
Comments: Moore (1859) described this
species under the genus Dasychira Stephens from northernIndia. Hampson
(1892) shifted it in the present genus and placed Gazalina venosata Walker
under it. This placement remained stable.
Phylogenetic Analysis
For the construction of the
phylogenetic tree related to known species of the genus Gazalina Walker,
the nucleotide sequences from the present study and the sequences retrieved
from the NCBI database were used (Table 1). The nucleotide sequences for the Gazalina
transversa (Moore) are not available in the NCBI database and thus not
included. Multiple sequence alignment was performed with CLUSTAL x software and
a phylogenetic tree was constructed using the maximum likelihood method (ML) in
MEGA (version 6) software. The high posterior probabilities depicted the
confidence of each branch in the phylogenetic tree. A confidence bootstrap
value of 100 was observed for Gazalina chrysolopha (MH795522.1); Gazalina
chrysolopha (HQ991385.1) and Gazalina apsara (KX863079.1).
The phylogenetic tree obtained from the nucleotide sequences belonging to
genera of Notodontidae family and subfamily Lymantriinae depicts that the genus
Gazalina Hübner has a close relationship with the two mentioned taxa and
thus has been taxonomically interchanging places between the two.
For strong validation of the
molecular analysis, three different phylogenetic trees were constructed using
three different genera, i.e., Phalera Hübner, Cerura Schrank, and
Clostera Samouelle of the family Notodontidae as out groups. The trees
(Figure 5–8) with outgroup as Phalera bucephala (MN696381) and Clostera
restitura (OR064768) provide a vague analysis as outgroups merge within
the clades formed by the other sequences. While the outgroup Cerura vinula
(MN696387) formed a separate clade with Epicoma melanosticta
(GU929763) having a significant bootstrap value of 100.
Discussion
Walker (1865) established this
genus with venosata Walker (apsara Moore) as its type species
from North-India under family Lymantriidae. The taxonomic position of genus Gazalina
Walker remained ambiguous due to its continuous reshuffling between subfamily
Lymantriidae of family Erebidae and family Notodontidae.
Gardner (1943) recorded secondary
setae on the mandibles of Thaumetopoea cheela Moore (Thaumetopoeinae)
and Godfrey (1984) noted the presence of two distinct mandibular setae in
Dudusinae. But in genus Gazalina Walker, the mandibular setae are
altogether absent (Gardner, 1943). The presence or absence of these setae can
also be utilized for stabilizing the systematic position of the present genus.
Kiriakoff (1970) considered the subfamily Thaumetopoeinae of processionary
moths as a self-standing family, i.e., Thaumetopoeidae. The rank of
Thaumetopoeinae, whether it is a subfamily or it receives a family status, had
been a long controversy till Miller (1911) concluded to give it the subfamily
rank. On the basis of cladistic analysis, he considered it as a homogeneous
clade within Notodontidae. Schintlmeister (2008) in his work to recognize
Thaumetopoeidae as a distinct family due to its probable link with
Lymantriidae. But, he did not give the basis on morphological cladistics or
molecular phylogeny to his notion. Zahiri et al. (2010) also recognized
this subfamily well nested within Notodontidae based on molecular phylogeny.
The subfamily Thaumetopoeinae is composed of approximately 100 species in 20
genera occurring in Africa (including Madagascar), the Mediterranean, Europe,
Asia, and Australasia in a belt from the Middle East to Taiwan, New Caledonia,
and Australia (Schintlmeister, 2013). Kobayashi & Nonaka (2016) analysed
the 28S ribosomal RNA genes to infer molecular phylogeny and recognized ten
subfamilies in the family Notodontidae. They discussed the genus Gazalina
Walker under subfamily Thaeumatopoeinae. On the basis of morphological and
molecular phylogenetic analyses of the group, Basso (2017) identified three
major clades in subfamily Thaumetopoeinae. The first clade includes the
Australian genera Epicoma Hübner and Ochrogaster Stephens and
possibly seven other still unexplored genera, for a total of 30 known species.
The second clade includes the African genera Anaphe Walker, Epanaphe Aurivillius,
Hypsoides Butler, Paradrallia Bethune-Baker, and five other genera,
for a total of 55 species. The third clade includes the African, Asian, and
European genera Gazalina Walker and Thaumetopoea Hübner for a
total of 18 species. The species within the third clade (Gazalina spp.
and Thaumetopoea spp.) are associated with economically important trees
such as Fagaceae (oaks, Quercus), Pinaceae (pines, Pinus) and Anacardiaceae
(pistachio, Pistacia) and can occasionally occur at high densities (outbreaks)
in the northern part of their range, such as the Mediterranean basin and Europe
for Thaumetopoea (Battisti et al. 2015) and the Himalaya foothills for Gazalina
(Rahman & Chaudhry 1992). Battisti et al. (2017) talked about the traits
which are shared by all species in this subfamily, most notably is the presence
of urticating setae on larvae, adults, or both. The eggs are laid in clusters,
and larvae are gregarious during the entire larval stage. In most species of
processionary moths, the larvae build silken tents, from which they forage for
food in a typical head-to-tail procession. This behaviour of moving in
procession has been documented for all the species of these three clades of
this subfamily.
The Co I sequence of Gazalina
chrysolopha (MH795522) and Gazalina apsara (MH790288) from the
present study were compared with the available reference sequences in Genbank
maintained by NCBI by using BLAST. The BLAST results showed 86% similarity of Gazalina
apsara with sequence (KX863079) from Pakistan and more than 96% similarity
of Gazalina chrysolopha with sequence (HQ991385) submitted by Ashfaq et
al. (2017) from Pakistan which signifies the geographic proximity. The
interesting fact is that these sequences also showed more proximity with
different genera of the subfamily Lymnatriinae and supported the ambiguous
placement of the genus Gazalina Walker under Notodontidae. For example,
the comparison of obtained sequences with sequences of species of three
different genera namely Epicoma Hübner, Axiocleta Turner, and Aglaossoma
Walker referable to the subfamily Thaumetopoeinae of the family Notodontidae
and three different genera, i.e., Artaxa Walker, Lymantria Hübner,
and Naroma Walker of the subfamily Lymantriinae, the similarity index
lies between 80–84 % with Thaumetopoeinae of the family Notodontidae and more
than 84% with the subfamily Lymantriinae of the family Erebidae (Table 5).
This suggested that the genus Gazalina
Walker can be placed under the subfamily Lymantriinae on the basis of
molecular analysis. As far as behavioral aspects are concerned, the adult moths
referable to the genus Gazalina Walker show no movements with
disturbance and feign dead similar to that of typical lymantrids. The adult
moth possesses distinct anal tuft which is a characteristic feature of adults
of subfamily Lymantriinae. The external morphological characters including
genitalic features completely conform to the characterization of subfamily
Lymantriinae except its wing venation which seems to be trifid. The internal
genitalic studies on the basis of bulbous constrictor muscular area, small and
transparent cuticular tube and its subapical entry position into aedeagus in
males and presence of oval lagena and origin of ductus seminalis from middle of
ductus bursae in females make both the studied species congeneric. However,
these can be differentiated on the basis of certain features such as shape of
testis, seminal vesicle-II, ductus ejaculatorius duplex, division of primary
simplex in males and shape of eggs, utriculus and length of accessory glands in
females. Though Gazalina possess a stable generic position, but its
placement in a proper family still remains a taxonomic mystery and accounts for
more studies and validation.
Conclusion
Walker (1865) established the
genus Gazalina Walker with venosata Walker (apsara Moore)
as its type species from northern India under family Lymantriidae. The
taxonomic position of this genus remained ambiguous due to its continuous
reshuffling between the subfamily Lymantriidae of the family Erebidae and the
family Notodontidae. Though the sequence analysis in present study showed the
proximity of its species with species of different genera of the subfamily
Lymnatriinae and supported the ambiguous placement of the genus Gazalina
Walker under Notodontidae. But on the basis of morphological, molecular,
larval, pupal and behavioural characteristics and thorough review of previous
works particularly of Kobayashi & Nonaka (2016) and Basso (2017), the genus
Gazalina has been placed under the subfamily Thaeumatopoeinae of
Notodontidae.
Table 1. Morphometry of internal
male genitalic organs of Gazalina chrysolopha (Kollar).
|
|
Organ |
Intraspecific range in length
(mm) |
Intraspecific range in width
(mm) |
|
1. |
Testis |
0.68–0.71 |
0.68–0.71 |
|
2. |
Seminal vesicle- I |
1.69–1.72 |
0.19–0.21 |
|
3. |
Seminal vesicle- II |
2.84–2.86 |
0.40–0.43 |
|
4. |
Vasa deferentia |
2.28–2.31 |
0.09–0.11 |
|
5. |
Ductus ejaculatorius duplex |
3.44–3.46 |
0.38–0.41 |
|
6. |
Accessory gland |
7.24–7.26 |
0.14–0.16 |
|
7. |
Primary simplex |
10.64–10.66 |
0.16–0.18 |
|
8. |
Constrictor muscular area |
1.09–1.11 |
0.25–0.28 |
|
9. |
Cuticular tube |
0.19–0.21 |
0.10–0.13 |
Table 2. Morphometry of internal
female genitalic organs of Gazalina chrysolopha (Kollar).
|
|
Organ |
Intraspecific range in length
(mm) |
Intraspecific range in width
(mm) |
|
1. |
Testis |
0.68–0.71 |
0.68–0.71 |
|
2. |
Seminal vesicle- I |
1.69–1.72 |
0.19–0.21 |
|
3. |
Seminal vesicle- II |
2.84–2.86 |
0.40–0.43 |
|
4. |
Vasa deferentia |
2.28–2.31 |
0.09–0.11 |
|
5. |
Ductus ejaculatorius duplex |
3.44–3.46 |
0.38–0.41 |
|
6. |
Accessory gland |
7.24–7.26 |
0.14–0.16 |
|
7. |
Primary simplex |
10.64–10.66 |
0.16–0.18 |
|
8. |
Constrictor muscular area |
1.09–1.11 |
0.25–0.28 |
|
9. |
Cuticular tube |
0.19–0.21 |
0.10–0.13 |
Table 3. Morphometry of internal
male genitalic organs of Gazalina apsara (Moore).
|
|
Organ |
Intraspecific range in length
(mm) |
Intraspecific range in width
(mm) |
|
1. |
Testis |
0.78–0.81 |
1.34–1.36 |
|
2. |
Seminal vesicle– I |
1.49–1.52 |
0.54–0.56 |
|
3. |
Seminal vesicle– II |
1.88–1.92 |
0.53–0.56 |
|
4. |
Vasa deferentia |
3.89–3.91 |
0.14–0.18 |
|
5. |
Ductus ejaculatorius duplex |
2.93–2.96 |
0.45–0.48 |
|
6. |
Accessory gland |
8.43–8.45 |
0.18–0.21 |
|
7. |
Primary simplex |
9.99–10.01 |
0.20–0.25 |
|
8. |
Constrictor muscular area |
0.48–0.51 |
0.23–0.26 |
|
9. |
Cuticular tube |
0.59–0.61 |
0.13–0.16 |
Table 4. Morphometry of internal
female genitalic organs of Gazalina apsara (Moore).
|
|
Organ |
Intraspecific range in length
(mm) |
Intraspecific range in width
(mm) |
|
1. |
Common terminal filament |
0.35–0.38 |
0.09–0.11 |
|
2. |
Egg tube |
13.83–13.86 |
0.64–0.66 |
|
3. |
Pedicel |
Absent |
Absent |
|
4. |
Lateral oviduct |
0.94–0.96 |
0.74–0.77 |
|
5. |
Common oviduct |
0.99–1.01 |
0.74–0.76 |
|
6. |
Spermathecal gland |
5.44–5.47 |
0.74–0.77 |
|
7. |
Spermathecal duct |
0.93–0.96 |
0.74–0.78 |
|
8. |
Utriculus |
0.88–0.91 |
0.35–0.38 |
|
9. |
Lagena |
0.09–0.11 |
0.08–0.11 |
|
10. |
Infundibulum |
0.09–0.12 |
0.74–0.76 |
|
11. |
Corpus bursae |
1.14–1.16 |
0.98–1.01 |
|
12. |
Ductus bursae |
1.18–1.21 |
0.64–0.66 |
|
13. |
Ostium bursae |
0.14–0.17 |
0.34–0.36 |
|
14. |
Ductus seminalis |
1.64–1.66 |
0.11–0.14 |
|
15. |
Bulla seminalis |
Absent |
Absent |
|
16. |
Accessory gland reservoir duct |
0.68–0.71 |
0.74–0.76 |
|
17. |
Accessory gland reservoir |
0.88–0.91 |
0.34–0.36 |
|
18. |
Accessory gland |
4.19–4.22 |
0.74–0.76 |
|
19. |
Vestibulum |
0.34–0.37 |
1.58–1.61 |
|
20. |
Vagina |
0.43–0.46 |
1.59–1.62 |
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