Journal of Threatened Taxa |
www.threatenedtaxa.org | 26 February 2021 | 13(2): 17712–17721
ISSN 0974-7907 (Online) | ISSN 0974-7893
(Print)
https://doi.org/10.11609/jott.6429.13.2.17712-17721
#6429 | Received 17 July 2020 | Final
received 15 September 2020 | Finally accepted 08 February 2021
Some biological aspects of the
central Indian endemic scorpion Hottentotta
jabalpurensis Kovařík,
2007 (Scorpiones: Buthidae)
Pragya Pandey 1, Pratyush P. Mohapatra 2 & D.B. Bastawade
3
1 University School of Environment
Management, Guru Gobind Singh Indraprastha
University, Dwarka Sector-16, New Delhi 110078, India.
2 Zoological Survey of India,
Central Zone Regional Centre, Vijay Nagar, Jabalpur, Madhya Pradesh 482002,
India.
3 Institute of Natural History
Education and Research (INHER), C26/9, Ketan Heights, Kothrud, Pune,
Maharashtra 411038, India.
1 pragya21.pp@gmail.com, 2 pratyush.m@zsi.gov.in
(corresponding author), 3 dbhushanbastawade@yahoo.co.in
Editor: Anonymity
requested. Date of
publication: 26 February 2021 (online & print)
Citation: Pandey, P., P.P. Mohapatra &
D.B. Bastawade (2021). Some biological aspects of the
central Indian endemic scorpion Hottentotta
jabalpurensis Kovařík,
2007 (Scorpiones: Buthidae) Journal of Threatened Taxa 13(2): 17712–17721. https://doi.org/10.11609/jott.6429.13.2.17712-17721
Copyright: © Pandey et al. 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: None.
Competing interests: The authors
declare no competing interests.
Author details: Pragya Pandey is a PhD scholar in the
University School of Environment Management, Guru Gobind
Singh Indraprastha University. She persued her MSc
dissertation on scorpions of Madhya Pradesh at Zoological Survey of India,
Jabalpur, Madhya Pradesh. Pratyush P. Mohapatra is Scientist at Zoological
Survey of India, Central Zone Regional Centre, Jabalpur, Madhya Pradesh. D.B. Bastawade is retired Scientist of Zoological Survey
of India and extensively studied Scorpions of India. He is continuing his
research by guiding students and educating researchers through the Institute of
Natural History Education and Research (INHER), Pune.
Author contribution: PP carried out field surveys,
reared the animals, conducted the research work and wrote the draft manuscript.
PPM supervised the research work, validated the information and helped in
manuscript writing. DBB validated the observations, guided the research work
and helped in editing the manuscript.
Acknowledgements: We thank Dr.
Kailash Chandra, Director, Zoological Survey of India and Dr.
S. Sambath, Scientist-E and officer-in-charge, ZSI,
Jabalpur for support and encouragement.
Thanks are also due to Mr. Manoj Dehury,
Vasundhara, Bhubaneswar, Odisha for preparation of the distribution map. We thank Saipari Sailo, Ashok K. Bhilala, Rajesh Somkuwar, R. Deepak, Rishabh Jaggi
and Deepeshwar Singh for help in field work and staff
of ZSI, CZRC, Jabalpur for overall administrative support.
Abstract: The present study reports the
distribution and bionomics of Hottentotta jabalpurensis Kovarik, 2007,
a central India endemic species.
Information on courtship, maternal care, feeding habits, and kin
recognition has been provided based on observations made in captivity. Morphological and morphometric details of the
pre-insemination spermatophore of the species have been detailed for the first
time for the species.
Keywords: Behaviour, breeding,
distribution, kin recognition, Madhya Pradesh, spermatophore.
Introduction
The genus Hottentotta
Birula, 1908 consisting of 55 species (Rein 2020)
is one of the most widely distributed genera of the family Buthidae,
distributed across Africa, the Arabian Peninsula, and in Asia (Kovařík 2007). In
India Hottentotta is represented by eight
species, namely: H. tamulus (Fabricius, 1798), H. pachyurus
(Pocock, 1897), H. rugiscutis (Pocock,
1897), H. jabalpurensis Kovařík, 2007, H. stockwelli
Kovařík, 2007, H. keralaensis
Ashwathi, Sureshan
& Lourenço, 2016, H. reddyi
Lourenço, 2015, and H. vinchu
Mirza, Ambekar & Kulkarni, 2019 (Kovařík 2007; Bastawade et al.
2012; Lourenço 2015, Ashwathi
et al. 2016; Mirza et al. 2009). The
genus is distributed in six out of 10 biogeographic zones of India (Rodgers et
al. 2000), namely Gangetic plains, desert, semi-arid, Deccan Peninsula,
Western Ghats, and coasts (Bastawade et al.
2012). Hottentotta
is a medically important genus (Ward et al. 2018) and has evolved to inhabit
closely to human dwellings, agricultural fields, and open areas (Tikader & Bastawade 1983; Ranawana et al. 2013; Mirza et al. 2019) which increases
its frequent interaction with humans. Hottentotta jabalpurensis
(Type locality: Jabalpur, Madhya Pradesh) was described based on some
morphological characters in which it differs from its sister species H. tamulus in having metasoma densely hirsute,
and the patella of pedipalp with long hair (Kovařík
2007).
Scorpions are predatory animals and are known for
their cannibalistic behaviour, however, apart from an obligatory mother-young
association, a few species are known to live in social groups. Sub-social behaviour has been observed in
species like Heterometrus fulvipes and H. swammerdemi
in the family Scorpionidae and on the other hand
cohabitation and gregarious interaction have been seen in Compsobuthus
werneri judaicus (Shivashankar 1994; Mohapatra & Pandey 2020; Warburg
2002). Similarly, studies on various
behavioural aspects of courtship and breeding have been undertaken on some
Indian scorpions of the family Scorpionidae and Buthidae (Bastawade 1992; Mirza
& Sanap 2009; Mirza et al. 2009; Mohapatra &
Pandey 2020).
Scorpions like many other arachnid groups have an
indirect way of sperm transfer through a sperm packet called spermatophore (Polis 1990). The courtship and breeding process can be
divided into four stages, i.e., initiation, Promenade á Deux, sperm transfer
and separation (Ross 2009). Furthermore,
spermatophore and hemispermatophores are considered
to have characters having taxonomic significance and phylogenetic values (Francke 1979; Monod et al. 2017). Among the Indian scorpions, studies on the
morphology of spermatophore and hemispermatophore are
relatively rare and are restricted to a few species (see Mathew 1956; Bastawade 1992, 1994; Mirza & Sanap
2009; Mohapatra & Pandey 2020).
Scorpions are viviparous arthropods with a long
gestation period (Polis 1990). The
females during parturition form a birth basket by using the first pair of legs,
crossing each other medially (Francke 1982). After birth, the babies climb up to the
mother’s back and settle in a particular orientation. The larval orientation can be random,
transverse, or longitudinal depending on different families or species (Savary 1996). They
continue to do so for a varying number of days after the young scorpions
undergo their first ecdysis to second-instar, after which the vagile
second-instar young ones disperse from the mothers’ dorsum and become
free-living (Williams 1971; Polis & Sissom 1990; Lourenço 2018).
Studies on such mother and young associations in Indian scorpion species
is very rare and has been mentioned by Mathew (1962) for Lychas
tricarinatus, Mirza et al. (2009) for Hottentotta pachyurus
(Pocock, 1897), and Mirza & Sanap (2009) for Heterometrus phipsoni
(Pocock, 1893).
The present study emphasizes the breeding behavior of H. jabalpurensis
along with distribution and natural history observations. Information on the
Morphological description of the spermatophore in the pre-insemination state,
parturition, maternal care, kinship behaviour and cannibalism has been provided
based on observations in captivity.
Materials and Methods
The study was undertaken during a seven months
dissertation work carried out by the first author from January to July 2018 at
the Zoological Survey of India (ZSI), Central Zone Regional Centre (CZRC), Jabalpur,
Madhya Pradesh. Scorpions were sampled
randomly by lifting rocks, finding them in leaf litter, peeling off bark and
digging the burrows in various sampling localities outside protected areas. During the night, scorpions were searched
with the aid of an ultraviolet torch.
Animals were handpicked with the help of a forceps when located and were
kept separately in plastic boxes to avoid being eaten by larger ones. Photographs were taken in their natural
habitat and the captive individuals were photographed regularly to record their
behavior with a Nikon D5100 camera fitted with
Nikon-100 macro or Tamron 90mm lens. A
total of 18 adult individuals of Hottentotta
jabalpurensis comprising four adult males and 14
adult females were collected from Sidh Baba Mandir
area, Katangi, and Paatbaba
area, Jabalpur, during the month of March–June 2018. All the animals except two males, from which
the spermatophores were obtained, were released back in their respective
habitats after the completion of the experiment during September 2019. To record various behavioral
aspects, live scorpions were maintained in the laboratory in terrariums or
plastic boxes of 6ˊˊX 3ˊˊX 3ˊˊ sizes, with a layer of 1ˊˊ soil substratum. The room temperature was maintained in an
air-conditioned room (24–30°C) and water was provided in each terrarium in
small bowls to maintain humidity and avoid desiccation. The animals were fed with live mealworms
maintained in the laboratory. Other prey
species such as gryllids, small geckos (Hemidactylus
spp.), skinks (Eutropis spp.), and termites
were also fed from time to time. Behavioral aspects such as feeding, courtship, and kin
recognition were recorded during the study period. Spermatophores of the scorpions were obtained
from the captive breeding groups and were preserved in 70% ethyl alcohol. Photographs were taken to see the natural
coloration. Morphological data were
collected under a stereo-zoom microscope (Leica M-2054) and measurements were
taken using Mitutoyo™ digital calipers to the nearest
0.1mm. Morphometric details of the
spermatophore were taken to the nearest 0.01mm under the microscope. Distribution localities of the species were
recorded based on the meta-data available for the specimens housed in the
national zoological collections of ZSI, CZRC.
Each specimen was identified by evaluating standard taxonomic characters
in a datasheet to record quantitative (i.e., mensural and meristic) and
qualitative taxonomic characters following standard taxonomic keys (Bastawade 1992; Kovařík 2007;
Lowe 2010; Monod et al. 2017).
Results
Distribution (Figure 1): Hottentotta jabalpurensis
was found to be distributed in Damoh, Sagar, Narsinghapur, Jabalpur, Chhindwara, Raisen, Dewas, Dheona, Panna, Shahdol, Shivpuri, Sidhi, Indore,
Hoshagabad, Ratlam, and Umaria districts of Madhya Pradesh (Table-1). This species was found mostly below rocks and
large boulders, sometimes clinging on to the inner surface of rocks and sitting
on wood logs in wet regions. Although,
this species was generally found solitary, males and females were found to be
under the same rock during January–March, probably due to breeding activities.
Feeding habits (Image 1,2): In captivity, scorpions were fed with
small crickets, mealworms, centipedes, termites and grasshoppers. Prey larger than the body size was stung to
overpower and feed on it. Juveniles
after a few days of second molting started using the
stinger and in comparison to adults the use of stinger was more frequent in the
juveniles. The juveniles were also found
to hunt in a group to overpower larger prey.
Adults as well as juveniles were found to manipulate the orientation of
the prey to feed on the head at first, no matter from which side it was
captured. The prey was eaten as a whole
or it was torn into pieces and the undigested parts were discarded after the full
meal. Smaller preys such as termites,
mealworms, etc., were devoured fully.
Sexual dimorphism: Marked sexual dimorphism was found in H. jabalpurensis.
Females were larger than males with a total length of 50–80 mm in
females and 48–65 mm in males. The males
also had a higher number of pectinal teeth (30–36)
than females (25–30). In males, the body
was slender with chela of pedipalp more robust than females and the males had a
prominent protuberance at the proximal end of the moveable chela and scalloped on
the immovable finger. Furthermore, adult
males were found to have yellowish legs, metasoma and pedipalps whereas the
females, were mostly reddish-brown overall.
Mating behavior (Image 3): The mating behaviour was observed in three
pairs and the findings are as follows.
The process took place in
captivity on three occasions on 06.iv.2018, 23.iv.2018, and 24.iv.2018
when the male and female were introduced into the same box. The courtship was always initiated by the
male and he started with patting the female and trying to grasp her from
pedipalps. Of the breeding pair on
06.iv.2018, the female remained immobile and withdrew the pedipalps after the
first attempt made by the male. The male
juddered several times while vigorously pulling the female towards him which
led to holding each other through pedipalps to perform the ritual dance
‘promenade á deux’.
Also, occasional cheliceral massage and tail
raising were observed. After a halt for
10–12 minutes, the male again pulled the female using just one pedipalp and
keeping away the unused one while constantly searching for a suitable
substratum with raised pectines. Occasionally the female resisted the
movements, but the male moved closer to her and tried pulling her with elevated
body and metasoma while the female kept her body close to the substratum with
raised pectines.
The male deposited the spermatophore on a small piece of wood (Image 6)
following vigorous juddering and waited for the spermatophore to get dried
up. The breeding process from initiation
till deposition of spermatophore lasted for 30 minutes. The second pair (on 23.iv.2018) bred in the
same way and the process lasted for 40 minutes.
The male deposited the spermatophore on a paper. Of the third pair, which was observed on
24.iv.2018, the courtship was initiated naturally when a male entered a
female’s compartment. The female in this
case did not resist the male, rather she was found juddering and trying to take
a hold of the male’s pedipalp. Although
juddering in this female was not as vigorous as it was observed in the males,
but the male did not produce spermatophore even after 45 minutes of courtship. Also, it was noted that in the above two
cases, the females did not show any interest in sperm transfer and the males
were always observed to escape the site immediately after mating. In another case where a male was kept with a
female, the pair did not show any affinity for courtship until 20 hours after staying
together. Another interesting behaviour
observed during the study period was when a male was found to be clasping the
female sitting very close to her and later fed upon the female after two days.
Spermatophore (Image 6–8):
Two pre-insemination spermatophores of H. jabalpurensis
bearing registration numbers ZSI-CZRC-7264 and 7265 were studied. The spermatophore when extruded was
semi-solid at the pedicle, which came out first and got firmly stuck on the
substratum. It was translucent with a
pinkish brown tinge, capsule dark brown, stem dark brown on the sides and paler
at the middle portion, flagellum whitish.
Pedicel flat, little broader at the base and creamy white. After extrusion, it turned solid and brownish
within five minutes. The capsular region
was reddish-brown comprising base, capsular distal carina, capsular basal
carina and basal hook. When the
spermatophore was extruded by the male it got glued to the substratum by the
pedicel at one end while the flagellum got attached to the other end of the
substratum. The stretching of
spermatophore probably helps in maintaining a particular direction after it
dries up.
The spermatophores (ZSI-CZRC-A-21455 and 21456) show
slight variation in stem length (6.4mm in the former and 6.7mm in the latter)
and length of the flagellum (4.9mm in the former and 3.7mm in the latter). The spermatophore of ZSI-CZRC-7264 is
described as follows. Most part of the flagelliform spermatophore comprises a tubular stem, which
is slender, elongated, translucent and hollow, 6.4mm in length, 0.9mm width and
0.7mm in depth. The capsule is the
complex part of the spermatophore and functions as a storage for spermatozoa
and responsible for sperm transfer. It
is broader at the base, 1.0mm in length and tapered towards the apex,
comprising a pair of tapered capsular distal carina (0.14mm in length), pair of
capsular basal carina, pair of stout and apically pointed basal hook (0.07mm
length). The capsule width at the region
of basal hook was 0.25mm. The flagellum
was 4.9mm long, extending from the ventral side of the capsular region and is
divided into a thicker part, pars recta, and a distal thinner part called pars reflecta. There is a
raised portion on the dorsal side of the flagellum just below the distal end of
the ventral process (termed as hook).
Parturition and maternal care (Image 4,5): Hottentotta
jabalpurensis females (n=9) gave birth to 23–45
juveniles in captivity. The parturition
mostly took place during early evening hours of 17.00–20.00 h. (n= 6), or during
late night 11.00–01.00 h. (n= 2) and on one occasion it took place at
14.00h. Prior to parturition the females
dug a pit using their first two pairs of her legs and was found resting in a
peculiar posture by bending its first pair of legs at the patellar-tibial joint
making a ‘birth basket’, thus fencing the area in front of the genital
operculum. During the process of
parturition, babies came out of the genital operculum at an interval of 3–5
minutes. After emergence, the babies
climbed onto the dorsum of the mother.
Babies were oriented randomly on the dorsum and were found stacked in
two or three layers on the dorsum, sometimes extended to the ventral part
also. It was observed that juveniles
remained on the dorsum till the completion of second molting
after which their exoskeleton becomes hardened and they start becoming
independent. It was observed that till
the second molt, the babies were not feeding,
although the mother was accepting food just after parturition. One among the female retained the parturition
after giving birth to five babies till the second ecdysis of the young ones
already on her body and later when it was shifted to another box after 10 days,
it again gave birth to 18 more babies.
Maternal care was found to be obligatory and while
carrying babies, the mother was very attentive, cautious and mostly
aggressive. In one incident one male was
introduced into the box having a gravid female and when it tried to interact
with the female, the female showed no interest at first but later got very
aggressive and attacked the male following which the male was separated
immediately. Juveniles became
independent and aggressive after the second molt,
i.e., after 8–10 days of birth but preferred to stay near the mother. While the babies were found moving away from
the mother, in the case of the slightest disturbance they immediately ran
towards the mother.
The difference between birth and first molt was 2–3 days and between the first and second molt this difference was 4–5 days (n=9). Scorplings were
pale yellow at birth with translucent and slight orange in color
pedipalps and metasoma, possessing clearly visible median and lateral carinae
on the mesosoma as well as dorsal, lateral and ventral carinae on the
metasoma. Chelicerae and legs are
translucent with black patches. They
have a soft exoskeleton and under-developed stinger. Hence, they
cannot sting or feed and utilize the stored nutrients. After the first molt
they start to change color from pale yellow to
brownish-orange, carinae becomes obtrusive, the exoskeleton gets thicker and
the stinger becomes hard and sharp.
Colour turns brownish-black on the mesosoma and legs and orange on the
pedipalp and metasoma with well-marked carinae after the second molt and characters become more prominent as they
grow. After the second molt, the babies started feeding on supplied prey and
sometimes, they consumed the leftover of the mother’s or other juvenile’s
prey. Cannibalism was observed to be
common among juveniles.
Kin recognition (Image 9): Juveniles interacted more than adults under
captivity where they either used to avoid or feed on each other. Also, sometimes mothers were found to feed on
babies soon after they were born. The
most interesting observation includes two juvenile bearing females coming in
contact and communicating with each other, being very close, patting one
another using pedipalps. The females
after being recognized stayed together for two days without harming each other
or any juveniles. No cannibalism was
found in this case and the juveniles of both the broods readily accepted
supplied prey. Later, after a few days
of second molting, when one of the females was kept
in another box, she gave birth to 18 more babies.
Discussion
Studies on Indian scorpions are mostly confined to
taxonomy and regional checklists; however, information on bionomics is still
understated. The present study is an
attempt to expand the information on biological aspects of a species based on
observations in captivity as well as their natural microhabitat. This central Indian endemic species, H.
jabalpurensis like other congeners such as H.
tamulus, H. pachyurus,
and H. vinchu, was found generally
below the rocks and large boulders, and sometimes on fallen wood logs in wet
regions. The coloration among males and
females and the juveniles could be added as intraspecific taxonomic characters
as the original description is based on old collections (Kovařík
2007). The present study extends the
distribution range of the species beyond its type locality in other parts of
Madhya Pradesh. As per the information
available on maternal care in H. pachyurus by
Mirza et al. (2009), the juveniles did not show much morphometric variation
from that of H. jabalpurensis; however,
the molting intervals differ as the former species
showed just 3–4 days’ time span between birth to the second molt
which is quite fast whereas it was found to be 8–10 days in the latter
species. Also, unlike H. pachyurus, juveniles in H. jabalpurensis were not found consuming their molt.
The spermatophore study on the Indian scorpion is
limited to a very few species, hence an attempt has been made to compare the
available data on spermatophore/ hemispermatophore of
genus Hottentotta. The spermatophore of H. tamulus as reported by Bastawade
(1992) varies by being placed at an angle of 20° and inverted backward whereas
the spermatophore of H. jabalpurensis
was placed at an angle of 10°. This
variation might be due to differences in the state of the spermatophore and the
inversion of H. tamulus spermatophore
could be due to the female exerting pressure on the capsule while obtaining the
spermatids as Bastawade (1992) described the
post-insemination spermatophore. As
mentioned by Bastawade (1992), the total length of
the spermatophore of H. tamulus is
1.3–1.5 times longer and the stem is twice the length than that of H. jabalpurensis (present study). Variations on the capsular region between
these two species could not be assessed because of a lack of mensural or meristic
data on the capsular structure for H. tamulus. Furthermore, in comparison with the
information on hemispermatophore of two chinese species, H. pellucidus
and H. saxinatans provided
by Lowe (2010) no comparative information could
be inferred.
Iteroparity is a common phenomenon in scorpions as
such reproductive strategy is observed in various species (de Albuquerquea & de Araujo Lira 2016) and there is some
information available on intervals between parturitions (Lees 1955; Mathew
1962; Polis & Farley 1979; Warburg 2012).
Among the Indian scorpions, Mathew (1962) discussed embryonic diapause
in Lychas tricarinatus
which was of 41–42 days. Our observation
in one of the Hottentotta jabalpurensis giving birth to babies within an interval
of 10 days is possibly a case related to unfavourable environmental
conditions. Another interesting behavior observed in this study is female juddering in
response to a male approaching for mating.
Juddering is a common phenomenon in males during mating as a direct response
to stimulate unreceptive females (Ross 2009).
Even during the present study, we observed juddering as a mode of
communication to suppress unreceptive counterparts. Furthermore, Hottentotta
jabalpurensis males holding the females by
engaging one of the pedipalps and keeping another free is an additional behavior recorded during the study. Hence, this study reports some additional
behavioural observations in Hottentotta jabalpurensis, which can also be studied in other
Indian species.
As envisaged from the study, there is no specific
threat to the species. This species is
highly adaptable and observed in various micro-habitat types ranging from
human-modified habitats to undisturbed forests.
H. jabalpurensis was found near
human habitation, agricultural fields, scrub forest, deciduous forest and
semi-evergreen forests and was found below small to large boulders, logs and
crevices. This species also occurs in a
large number of protected areas such as Veerangana Durgavati WS, Singhori WS, Kanha TR, Bandhavgarh TR, Satpura TR, Bori WS, and Sanjay
NP. As this species is considered potentially dangerous for humans and its
envenomation has the possibility of confusion with snake-bite, study on
human-scorpion interaction can be undertaken to understand the prevalence of
such conflicts. A case of envenomation
as observed by one of the authors (PPM), following the sting on the right-hand
thumb is described as follows. There was
swelling and systemic pain around the affected area with a feeling of dizziness,
followed by uncoordinated movements for about 6hr. The swelling became normal after 56–60 h
following the envenomation.
Table 1. Geocoordinates of the localities of
distribution of Hottentotta jabalpurensis in Madhya Pradesh (WGS-84).
|
Location |
GPS
coordinates |
|
Damoh |
N23.8210°,
E79.4514° |
|
Narsinghpur |
N22.9473°,
E79.1923° |
|
Sagar |
N23.8388°,
E78.7378° |
|
Ranjhi,
Jabalpur |
N23.1815°,
E79.9864° |
|
Patbaba
area, Jabalpur |
N23.1702°,
E79.9747° |
|
Sidh
Baba Mandir area, Katangi |
N23.4645°,
E79.7980° |
|
Nauradehi
WLS |
N23.3683°,
E79.1718° |
|
Chhindwara |
N22.0574°,
E78.9382° |
|
Barna
reservoir, Raisen |
N23.0795°,
E78.0310° |
|
Kartholi, Dewas |
N32.6307°,
E74.9490° |
|
Veerangana Durgavati WLS, Damoh |
N21.6970°,
E77.7954° |
|
Kesli, Sagar |
N23.4226°,
E78.8184° |
|
Madhav NP,
Shivpuri |
N25.4317°,
E77.7391° |
|
Lameta ghat, Jabalpur |
N23.1092°,
E79.8282° |
|
Kala Dehi, Jabalpur |
N22.8787°,
E79.8293° |
|
Panna TR,
Panna District |
N24.7166,
E80.2000 |
|
Shahdol |
N23.3022,
E81.3267° |
|
Sidhi |
N24.3956°,
E81.8825° |
|
Indore |
N22.6791°,
E75.8580° |
|
Singhori
WLS, Kheoni |
N22.1983°,
E79.0579° |
|
Pachmarhi BR |
N22.4674°,
E78.4346° |
|
Ratlam |
N23.3315°,
E75.0367° |
|
Bandhavgarh N.
Park, Umaria |
N23.7224°,
E81.0242° |
For figure
& images - - click here
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