Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October
2019 | 11(13): 14672–14680
Moulting pattern and mortality
during the final emergence of the Coromandel Marsh Dart Damselfly
Ceriagrion coromandelianum
(Zygoptera: Coenagrionidae)
in central India
Nilesh R. Thaokar
1, Payal R. Verma
2 & Raymond J. Andrew 3
1–3 Centre
for Higher Learning and Research in Zoology, Hislop College, Civil lines,
Nagpur, Maharashtra 440001, India.
1 nilesh.thavkar@gmail.com,
2 payalrverma@gmail.com, 3 rajuandrew@yahoo.com
(corresponding author)
doi: https://doi.org/10.11609/jott.4954.11.13.14672-14680
Editor: K.A. Subramanian, Zoological Survey of India, Chennai,
India. Date of publication: 26 October
2019 (online & print)
Manuscript details: #4954 | Received 15 March 2019 |
Final received 12 June 2019 | Finally accepted 02 October 2019
Citation: Thaokar, N.R., P.R. Verma
& R.J. Andrew (2019). Moulting pattern and mortality during the final
emergence of the Coromandel Marsh Dart Damselfly Ceriagrion
coromandelianum (Zygoptera:
Coenagrionidae) in central India. Journal of Threatened Taxa 11(13): 14672–14680; https://doi.org/10.11609/jott.4954.11.13.14672-14680
Copyright: © Thaokar et al. 2019. Creative Commons Attribution
4.0 International License. JoTT allows unrestricted use, reproduction, and
distribution of this article in any medium by adequate credit to the author(s)
and the source of publication.
Funding: Self funded.
Competing interests: The authors declare no competing
interests.
Author details: Dr. Nilesh R. Thaokar is presently teaching at the Post Graduate Department
of Zoology as an assistant lecturer. Dr. Payal R. Verma is actively engaged in research in the area of
odonatology.
She has completed her PhD from Rashtrasant Tukadoji Maharaj Nagpur University, and currently is an
assistant professor lecturer at the Post Graduate Department of Zoology. Dr. R.J.
Andrew has been studying various physiological, morphological,
ethological, and ecological aspects of dragonflies of central India for the
last 30 years and has more than 100 research papers to his credit. He serves as the director of the P.G. Dept.
of Zoology, and Vice Principal, Hislop College, Nagpur. He has published two books on odonates and has organized one international, three
southern Asian, six national, and two state level symposia.
Author contribution: NRT and PRV contributed in field work and documentation of the oviposition behaviour. RJA set
up the project and evaluated the findings.
Acknowledgements: We thank the Principal Dr.
Ms. Dipti R. Christian and Management of Hislop College, Nagpur for providing
us laboratory facilities.
Abstract: The final emergence of the Coromandel Marsh Dart
Damselfly Ceriagrion coromandelianum
was studied for 50 days (22 January–12 March, 2011) from the botanical garden
of Hislop College, Nagpur, India, (a semi controlled site) where small
underground cement tubs/tanks are used to grow macrophytes by the Botany
department. In C. coromandelianum
emergence is asynchronous, diurnal and occurs between 07.00h and 18.00h. Stage-I starts when the ultimate instar nymph
of C. coromandelianum leaves the water body,
searches for a suitable place and then begins to shudder its body to detach the
trapped pharate from the nymphal exuvia. The pharate exerts pressure on the thoracic
tergites to split the cuticle. Stage-II
starts when the head and thorax of the pharate emerges out of the split exuvia. The pharate
struggles to remove its trapped body from the nymphal
exuvia. During
Stage-III, the wings expand but are opaque; pigmentation of the body occurs
simultaneously all over the body. Soon
the whole body develops its species specific coloration while the expanding
wings gain transparency, unfold and separate out and now the imago is ready for
its maiden flight. Stages I, II, and III
occupy 31.66%, 11.73%, and 56.60% of the total moulting period,
respectively. A total of 243 emergences
occurred during the observation period, 158 emergences occurred in tanks
containing Pistia stratiotes, while 65
emergences in tubs containing Nymphaea nouchali
indicating that C. coromandelianum prefers P.
stratiotes over N. nouchali for
oviposition. Twenty deaths were recorded
during the present observation. Failure
to moult (15%) and failure to emerge completely out of the exuvia
(85%) were the two reasons for mortality.
Keywords: Dragonfly, emergence, exuvia,
instar, metamorphosis, moulting, pharate.
INTRODUCTION
In Odonata, moulting during the
final emergence when the aerial imago is released from the exuvia
of aquatic nymph is a fascinating event involving many different types of
rhythmic movements. It is also a very
vulnerable period since the helpless individual is exposed to various
antagonistic factors of the environment.
This process was documented by various workers (Tillyard
1917; Corbet 1957; Pajunen 1962; Trottier 1966; Ubukata 1981; Banks & Thompson 1985; Gribbin & Thompson 1990, 1991; Haslam, 2004; Andrew
& Patankar 2010) and was evaluated by Corbet
(1999) who divided it into four observable stages. Later, Andrew & Patankar
(2010) modified this division and proposed only three stages taking into
consideration the time-lag and attainment of morphological characters of the
freshly moulted imago. Eda (1963),
reported two major types of posture during emergence- horizontal emergence
commonly found in Zygoptera and Gomphidae
and the vertical, found in the remaining groups, though inverted emergence has
also been reported in some species of Zygoptera (Rowe
1987). Mortality during emergence can be
caused by three observable factors: failure to moult, failure to harden
body/wings, and predation (Thompson 1991; Bennett & Mill 1993; Andrew
2010). It can range from 0% to 100% and
is dependent upon factors like temperature, rain, wind, oxygen level, lack of
suitable emergence support, overcrowding and predation (Corbet 1957, 1999; Pajunen 1962; Kurata 1974; Inoue
1979; Thompson et al. 1985; Gribbin & Thomas
1990; Bennett & Mill 1993; Jacob & Suhling
1999; Purse & Thompson 2003; Andrew 2010).
Most studies on the final emergence in Odonata are confined to species
of the sub-tropical and temperate regions, while only a few attempts have been
undertaken to study this process in detail in the tropical region mostly
covering only the anisopteran species (Mathavan & Pandian 1977; Andrew 2010, 2012; Andrew
& Patankar 2010).
The zygopteran
Ceriagrion coromandelianum
(Fabricius, 1798) is a very common damselfly of the
Indian subcontinent. The life history of
this species was described in detail by Kumar (1980) and Sharma (2009). Kumar (1980) also described the larval
morphology of all the instars. We have
used this species to evaluate various aspects of odonate
reproductive biology (Andrew et al. 2011a,b; Thaokar
et al. 2018a,b). It is found almost
throughout the year, ovipositing in various floating
and submerged vegetations of small natural and man-made water bodies (Sharma
2009; Andrew et al. 2011a). The present
paper describes the pattern and process of emergence of this damselfly with a
note on mortality during this event.
MATERIAL
AND METHODS
Site: The observation was carried out
at the botanical garden of Hislop College, Nagpur, (21.1470’N,
79.0710E), India, a semi-controlled site, where small underground
cement tubs are used to grow macrophytes by the Botany department. The tubs contain floating Nymphaea nouchali, Lemna paucicostata and submerged Hydrilla verticillata vegetation, while the cement tank contains
only Pistia stratiotes. These are surrounded by bushes of
flowering plants and post-noon, this area is under the shadow of the college
building. Ceriagrion
coromandelianum is found breeding all round the year at this site.
Mature F-0 larvae were collected
from this site and kept in a glass tank partially filled with water along with
floating vegetation. Natural conditions
were maintained by keeping the containers near the large open windows. With the
help of an aim-n-shoot Sony (DSC-W30) and Canon (G11) cameras, various stages
of the process of moulting during metamorphosis were documented. All movements of the larva/emerging pharate were
documented and an electronic stopwatch was used to record the time. Some of the emergences were directly recorded
at the collection site. Daily collection
of exuviae was undertaken at the study site from 22
January to 12 March, 2011 (Table 1).
Details of the weather report of the city were procured from the website
https://www.timeanddate.com.
RESULTS
The daily emergence of Ceriagrion coromandelianum
was recorded for 50 days by collecting the exuviae
from the water tubs of the above described site from 22 January to 12 March
2011. The tubs and tank were filled with
floating Pistia stratiotes and Nymphaea
nouchali which made a perfect substrate for the
final emergence. A total of 243 emergences occurred during the observation
period of 50 days at the study site (excluding the ones collected from the
study site and reared in the laboratory), 158 emergences occurred in tanks
containing Pistia stratiotes while 65
emergences in tanks containing Nymphaea nouchali. Two peaks of emergences were recorded, the
first emergence on 14 February (19) and the second on 21 February (17) (Table
1). Fifty percent of the total emergence
of C. coromandelianum was observed by the 29th
day (19 January 2011) (Fig. 1). The
duration of the day was divided as morning (07.00–12.00 h), noon (12.00–16.00
h), and evening (16.00h–dusk). Emergence
was not found during the pre-dusk and pre-dawn period. The number of emergence recorded were:
morning 58 (23.86%), noon 166 (68.31%), and evening 19 (7.8%) (Table 2, Figure
2a). The highest number of emergence
(22) was observed on 14 February 2011 (Max. temp. 34oC, min. temp 20oC,
humidity 30% at noon) followed by 18 emergence on 20 February 2011 (Max. temp.
25oC, min. temp. 19oC, humidity 85% at noon) (Figure
2b). Depending upon the type of
substrate C. coromandelianum can moult in both
horizontal (on floating leaves) as well as vertical (on emerging stem)
positions. Eleven complete events of
metamorphosis leading to emergence of the pharate were observed and recorded. Moulting in C. coromandelianum
is not time specific since this process occurs throughout the day between
07.00h and 18.00h (Table 3).
The following documentation
describes one complete pattern of moulting during the final emergence of the
damselfly, C. coromandelianum observed on 18
February 2011, which started at 12.55h and ended at 15.38h, (153 minutes)
(Images 1–9). This process has been
divided into three observable stages (Andrew & Patankar
2010).
Stage- I: At 12.55h the F–0 larva emerged
out of water and climbed the floating leaf of Pistia
stratiotes. It moved 4cm on the dry
surface of the leaf, and rested. At
13.24h the larva began to shake the abdomen in the vertical plane. These movements were very slow and later it started
moving it in the horizontal plane. This movement continued for 56sec. Then it started pushing the head and thorax
against the leaf. The legs were spread
while the posterior region of the abdomen was firmly pressed against the
base. The larva moved the head sideways
and curved up the abdominal tip. It
reset the grip of the fore and hind legs and raised the head and thorax. This movement continued interspaced with long
intervals of motionless rest. It flexed
the legs to elevate the anterior region of the body. At 14.03h, a split appeared along the cuticle
of the dorsal region of thorax. This
concluded Stage-I of moulting which took 68 minutes.
Stage- II: Within two minutes the head and
thorax just elevated from the split exuvia without
wriggling, leaving the exuvia on the leaf. The legs were straight sticking along the
dorsal side of the body. At 14.07h, half the abdomen along with the head and
thorax was outside the exuvia. The legs started flexing slowly. Initially only the forelegs exhibited
movement but by 14.09h all the legs started moving and pawing the air without
touching the substrate (leaf). The body
of the pharate was still supported by the trapped abdomen. The thorax and abdomen formed an angle of 90
degree. The tiny compact wings lay
parallel to the abdomen. The pharate now
started making feeble movement of the legs trying to grip the leaf
surface. As soon as it found a suitable
grip for all the legs, the pharate smoothly extracted the remaining part of the
abdomen from the exuvia without wriggling. It was 14.14h and the end of Stage-II. This stage took only 11 minutes.
Stage- III: The fore and mid legs of the
pharate rested on the leaf while the hind legs now rested on the exuvia. It swayed
the body forward and straightened the curved abdomen and swayed back to the
original position without moving the legs.
Slowly, the telescoped abdomen started expanding. Concomitantly, the wings also started stretching
and by 14.42h, the wings were completely stretched but opaque white in colour
and still stuck to each other. The
pharate was motionless just re-adjusting the legs and re-gripping the leaf at
regular intervals. The abdomen continued
to expand and by 15.08h it was completely stretched and stiff. While the abdomen was expanding the pharate
cleared the gut by forcefully expelling water (23 times) from the rectum at
regular intervals. But for the eyes and
a slight tinge of green on the thorax, the pharate was un-pigmented (at this
stage, pink inter-segmental bands are observed in females which dissipate
within a few minutes). Pigmentation of
the body took place simultaneously all over the surface along with transparency
of the wings and by 15.27h the freshly emerged imago became flight worthy. The imago now exhibited its characteristic
species specific color patterning on the adult
body. Stage-III took 74 minutes.
A comparative account of 11
complete metamorphoses on site and in the laboratory shows that on an average,
the duration of Stage-I is 31.5 minutes (29.35%), Stage-II is 14 minutes
(13.04%), and Stage-III is 61.83 (57.61%) in the laboratory. While on the site the average duration of
Stage-I is 348 minutes (333.80%), Stage-II is 15 minutes (10.50 %), and in
Stage-III it is 79.2 minutes (55.70%).
Further, the average time to complete emergence is much higher on site
(142.5 minutes) as compared to in the laboratory (107.33 minutes) (Tables 3,
4).
The mortality rate recorded
during emergence was 8.2% (N= 20).
Failure to moult (15%, Stage-I) and the failure to emerge out of the exuvia (85%, Stage-II) were the two reasons of
mortality. During Stage-II, if the
pharate is unable to extract the abdomen and wings from the exuvia
within the optimal period, it results into a deformed imago (which may step out
of the exuvia) with twisted, telescoped abdomen and
crumpled, deformed wings ultimately
leading to the death of the individual (Images 10–13).
Case of an unsuccessful emergence
On 18 February 2011, one larva
was found out of water, preparing for the final emergence. By 12.48h a split was observed on the thorax
and slowly the pharate extracted the head and thorax from the exuvia. Half the way
it stopped and after a gap of three minutes it again started pulling itself out
of the exuvia.
At 12.58h, the wing buds along with the cuticle of the exuvia partly separated from the main body of the exuvia. The pharate
struggled to pull out the wings from the exuvia wing
bud case but with little success. Soon
the body of the phatrate was completely out of the exuvia along with a major portion of the wings but the wing
tips were still trapped. At 13.10h the
part of the wings outside the exuvia started
stretching and spreading and soon turned transparent, but the pharate could not
release the trapped wing tips. By 14.13h
although the complete body stretched but the wings lay trapped in the cuticle
resulting in an adult with deformed wings (Images 14–19).
DISCUSSION
There are two basic type of
postures during emergence, the upright type where the larva completes its
moulting at 0o between body and exuvia as
found in most Coenagrionidae, Gomphidae,
Lestidae, Petaluridae, and
the hanging type found in Aeshnidae, Calopterygidae, Corduliidae, and Libellulidae where the larva completes its moulting at an
angle which ranges from 90 to 130 degrees and therefore it becomes necessary
for the hanging type to climb on a vertical support substrate (Inoue 1964;
Trottier 1966). Although horizontal
emergence is common in Zygoptera and Gomphidae, inverted emergence occurs in the zygopteran Xanthocnemis
sinclairi and some Ischnura
spp. (Row 1987; Corbet 1999). Libellulidae mostly moult in a vertical position (Andrew
2010, 2012; Andrew & Patankar 2010). When the angle of emergence is manipulated,
the larva tries to regain its original positioning by readjusting its body (Calopteryx,
Heymer 1972) or by darting towards a vertical
substrate (P. flavescens, Andrew & Patankar 2010). C.
coromandelianum appears to be an opportunistic
species and can moult in both horizontal as well as vertical position. In C. coromandelianum,
the spreading of the wings is uniform as found in most libellulids
but in most horizontal emergence the spreading starts from the base upwards
(Eda 1963). Further in the hanging type
of emergence, gravitational force plays an important role in setting the angle
of the spreading wings with respect to the linear position of the body (Andrew
& Patankar 2010), but in C. coromandelianum gravity does not have any
influence as it exhibits both vertical and horizontal emergence. In P. flavescens,
the pigmentation of the body starts from the thorax and terminal end of the
abdomen (Andrew & Patankar 2010) but in C. coromandelianum pigmentation of the body takes place
all over the surface, simultaneously.
Variation in the number of
emergences during morning, afternoon and evening indicate that photoperiod and
temperature have a direct bearing on the initiation of emergence in C. coromandelianum. Purse
& Thompson (2003) reported that emergence in the damselfly Coenagrion mercuriale
was positively correlated to the duration of sunlight of the previous day. Positive correlation was found between
sunlight and daily emergence in Lestes eurinus (Lutz 1968).
In C. coromandelianum too, maximum
emergence is noticed during the afternoon period indicating a link between
intensity of sunlight on emergence, but no statistically significant
relationship could be established between daily temperature and humidity. Farkas et al. (2012) reported that in gomphid
dragonflies, inter year variations found during emergence is due to annual
fluctuation in the water temperature which may influence onset and synchrony of
emergence. A comparative account
of the time lag between the three stages of emergence between the anisopteran P. flavescens
and zygopteran C. coromandelianum
indicates that the major time of emergence is consumed in Stage-III for the
stretching and spreading of the body and wings in both the groups.
Mortality during emergence is
classified into three observable events: failure to moult, failure to expand
& harden the wings, and predation.
The first two are caused by factors such as low temperature, rain, wind,
low oxygen level, lack of suitable emergence support and overcrowding (Corbet
1999). Lack of mass emergence results in
little competition for support and eliminates overcrowding as a cause of
mortality in Onychogomphus uncatus and Orthetrum
coerulescens (Jakob & Suhlingg
1999) and a similar situation is also found in the observation. In the northern range margins of Britain,
Purse & Thompson (2003) reported a low mortality rate of 4.9% including
deformed individuals during the emergence of the damselfly Coenagrion
mercuriale. In
the damselfly Pyrrhosoma nymphula in Yorkshire, England the mortality during
emergence ranged 3–5% and was mainly due to incomplete ecdysis, failure to
expand wings and predation by spiders (Bennett & Mill 1993). In southern India, Mathavan
& Pandian (1977) reported that the mortality rate of most libellulid dragonflies varied between 8% and 14% during
emergence, whereas in central India the mortality rate of P. flavescens was 10.93% (Andrew 2010). In the Indian subcontinent predation rate is
very less, and ranges from zero to 0.78% (Mathavan
& Pandian 1977; Andrew 2010, 2012).
Failure to moult at Stage-I indicates that there may be some endogenous
(genetic) factors or injuries or dehydration which can be responsible for
mortality at this stage, whereas failure to emerge out of the exuvia occurs in Stage-II and could be caused by loss of
energy during moulting or difficulty in removing the trapped abdomen or wings
from the exuvia (Jakob & Suhling
1999; Andrew & Patankar 2010). Strong winds are a major cause of mortality
in Stage-III (Corbet 1999). In P. flavescens, 56% of the total mortality was found in
Stage-III at an open drain in central India (Andrew 2010). In this study, we did not observe a single
case of mortality at Stage-III probably because the site is well sheltered
against strong winds by the surrounding building of the institution. In the present study, predation was not
observed probably due to a lack of major predators at the semi controlled study
site. Further, it couldn’t be ignored
that mortality will be more in natural habitats in and around areas where
nesting density of predatory birds is high or where the pharate is more exposed
to extreme physical factors (Corbet 1999).
Jakob & Suhling (1999) found that the
predatory rate during moulting in odonates is mostly
less than one in most natural conditions.
Nevertheless, we (Thaokar et al. 2018a,b)
earlier reported that C. coromandelianum displays
a refined hierarchy of preferences for oviposition and chooses floating leaves
of Nymphaea nouchali over Lemna
paucicostata and submerged Hydrilla verticillata but with the addition of Pistia stratiotes at the site, C. coromandelianum prefers P. stratiotes over N.
nouchali for oviposition.
Table 1. Ceriagrion
coromandelianum: Number of emergence and
mortality observed during the 50-day study period (mortality in parenthesis).
|
|
Date |
Pistia stratiotes |
Nymphaea nouchali |
Mortality |
Total |
|
1 |
22.i.2011 |
0 |
0 |
0 |
0 |
|
2 |
23.i.2011 |
2 |
0 |
0 |
2 |
|
3 |
24.i.2011 |
1 |
0 |
0 |
1 |
|
4 |
25.i.2011 |
2+(1) |
0 |
1 |
3 |
|
5 |
26.i.2011 |
1 |
0 |
0 |
1 |
|
6 |
27.i.2011 |
3+(1) |
0 |
1 |
4 |
|
7 |
28.i.2011 |
0 |
0 |
0 |
0 |
|
8 |
29.i.2011 |
0 |
1 |
0 |
1 |
|
9 |
30.i.2011 |
2 |
2+(1) |
1 |
5 |
|
10 |
31.i.2011 |
1 |
0 |
0 |
1 |
|
11 |
01.ii.2011 |
0 |
1 |
0 |
1 |
|
12 |
02.ii.2011 |
0 |
0 |
0 |
0 |
|
13 |
03.ii.2011 |
1 |
2+(1) |
1 |
4 |
|
14 |
04.ii.2011 |
0 |
0 |
0 |
0 |
|
15 |
05.ii.2011 |
3 |
1 |
0 |
4 |
|
16 |
06.ii.2011 |
5+(1) |
1 |
1 |
7 |
|
17 |
07.ii.2011 |
2 |
0 |
0 |
2 |
|
18 |
08.ii.2011 |
2 |
0 |
0 |
2 |
|
19 |
09.ii.2011 |
1 |
2 |
0 |
3 |
|
20 |
10.ii.2011 |
1+(1) |
0 |
1 |
2 |
|
21 |
11.ii.2011 |
2 |
0 |
0 |
2 |
|
22 |
12.ii.2011 |
4 |
0 |
0 |
4 |
|
23 |
13.ii.2011 |
12 |
4 |
0 |
16 |
|
24 |
14.ii.2011 |
16+(3) |
3 |
3 |
22 |
|
25 |
15.ii.2011 |
6 |
3 |
0 |
9 |
|
26 |
16.ii.2011 |
2 |
3 |
0 |
5 |
|
27 |
17.ii.2011 |
4 |
1 |
0 |
5 |
|
28 |
18.ii.2011 |
3+(1) |
2+(1) |
2 |
7 |
|
29 |
19.ii.2011 |
6 |
0 |
0 |
6 |
|
30 |
20.ii.2011 |
12+(1) |
5 |
1 |
18 |
|
31 |
21.ii.2011 |
8 |
5 |
0 |
13 |
|
32 |
22.ii.2011 |
10+(2) |
1 |
2 |
13 |
|
33 |
23.ii.2011 |
4 |
5 |
0 |
9 |
|
34 |
24.ii.2011 |
5+(2) |
1 |
2 |
8 |
|
35 |
25.ii.2011 |
5 |
3 |
0 |
8 |
|
36 |
26.ii.2011 |
0 |
0 |
0 |
0 |
|
37 |
27.ii.2011 |
2 |
4+(1) |
1 |
7 |
|
38 |
28.ii.2011 |
4 |
1 |
0 |
5 |
|
39 |
01.iii.2011 |
5 |
2 |
0 |
7 |
|
40 |
02.iii.2011 |
0 |
0 |
0 |
0 |
|
41 |
03.iii.2011 |
0 |
0 |
0 |
0 |
|
42 |
04.iii.2011 |
2 |
2 |
0 |
4 |
|
43 |
05.iii.2011 |
5+(1) |
1 |
1 |
7 |
|
44 |
06.iii.2011 |
4 |
2 |
0 |
6 |
|
45 |
07.iii.2011 |
1 |
2 |
0 |
3 |
|
46 |
08.iii.2011 |
3 |
0 |
0 |
3 |
|
47 |
09.iii.2011 |
5+(1) |
3+(1) |
2 |
10 |
|
48 |
10.iii.2011 |
1 |
0 |
0 |
1 |
|
49 |
11.iii.2011 |
0 |
0 |
0 |
0 |
|
50 |
12.iii.2011 |
0 |
0 |
0 |
0 |
|
|
Total |
158+(15) |
63+(5) |
20 |
241 |
Table 2.
Ceriagrion coromandelianum:
Number of emergence at different period of the day.
|
|
Date |
Morning |
Afternoon |
Evening |
Total |
|
1 |
22.i.2011 |
0 |
0 |
0 |
0 |
|
2 |
23.i.2011 |
0 |
2 |
0 |
2 |
|
3 |
24.i.2011 |
0 |
1 |
0 |
1 |
|
4 |
25.i.2011 |
0 |
3 |
0 |
3 |
|
5 |
26.i.2011 |
0 |
1 |
0 |
1 |
|
6 |
27.i.2011 |
1 |
3 |
0 |
4 |
|
7 |
28.i.2011 |
0 |
0 |
0 |
0 |
|
8 |
29-.i.2011 |
0 |
1 |
0 |
1 |
|
9 |
30.i.2011 |
2 |
3 |
0 |
5 |
|
10 |
31.i.2011 |
0 |
1 |
0 |
1 |
|
11 |
01.ii.2011 |
0 |
1 |
0 |
1 |
|
12 |
02.ii.2011 |
0 |
0 |
0 |
0 |
|
13 |
03.ii.2011 |
2 |
2 |
0 |
4 |
|
14 |
04.ii.2011 |
0 |
0 |
0 |
0 |
|
15 |
05.ii.2011 |
1 |
2 |
1 |
4 |
|
16 |
06.ii.2011 |
2 |
4 |
1 |
7 |
|
17 |
07.ii.2011 |
0 |
2 |
0 |
2 |
|
18 |
08.ii.2011 |
0 |
2 |
0 |
2 |
|
19 |
09.ii.2011 |
1 |
2 |
0 |
3 |
|
20 |
10.ii.2011 |
0 |
2 |
0 |
2 |
|
21 |
11.ii.2011 |
1 |
1 |
0 |
2 |
|
22 |
12.ii.2011 |
1 |
2 |
1 |
4 |
|
23 |
13.ii.2011 |
3 |
11 |
2 |
16 |
|
24 |
14.ii.2011 |
6 |
13 |
3 |
22 |
|
25 |
15.ii.2011 |
2 |
6 |
1 |
9 |
|
26 |
16.ii.2011 |
1 |
4 |
0 |
5 |
|
27 |
17.ii.2011 |
1 |
3 |
1 |
5 |
|
28 |
18.ii.2011 |
2 |
5 |
0 |
7 |
|
29 |
19.ii.2011 |
3 |
3 |
0 |
6 |
|
30 |
20.ii.2011 |
4 |
11 |
3 |
18 |
|
31 |
21.ii.2011 |
4 |
8 |
1 |
13 |
|
32 |
22.ii.2011 |
5 |
7 |
1 |
13 |
|
33 |
23.ii.2011 |
1 |
8 |
0 |
9 |
|
34 |
24.ii.2011 |
3 |
5 |
0 |
8 |
|
35 |
25.ii.2011 |
3 |
4 |
1 |
8 |
|
36 |
26.ii.2011 |
0 |
0 |
0 |
0 |
|
37 |
27.ii.2011 |
2 |
5 |
0 |
7 |
|
38 |
28.ii.2011 |
2 |
3 |
0 |
5 |
|
39 |
01.iii.2011 |
2 |
4 |
1 |
7 |
|
40 |
02.iii.2011 |
0 |
0 |
0 |
0 |
|
41 |
03.iii.2011 |
0 |
0 |
0 |
0 |
|
42 |
04.iii.2011 |
0 |
4 |
0 |
4 |
|
43 |
05.iii.2011 |
1 |
6 |
0 |
7 |
|
44 |
06.iii.2011 |
1 |
5 |
0 |
6 |
|
45 |
07.iii.2011 |
0 |
3 |
0 |
3 |
|
46 |
08.iii.2011 |
0 |
3 |
0 |
3 |
|
47 |
09.iii.2011 |
1 |
8 |
1 |
10 |
|
48 |
10.iii.2011 |
0 |
1 |
0 |
1 |
|
49 |
11.iii.2011 |
0 |
0 |
0 |
0 |
|
50 |
12.iii.2011 |
0 |
0 |
0 |
0 |
|
|
Total |
58 |
165 |
18 |
241 |
Table 3. Ceriagrion coromandelianum—duration and average timing (in
minutes) of the three stages of the final emergence recorded in the laboratory.
|
Time (h) |
Stage I |
Stage II |
Stage III |
Total |
|
12.37–14.45 |
36 |
14 |
78 |
128 |
|
13.23–14.56 |
30 |
15 |
48 |
93 |
|
14.14–14.44 |
20 |
8 |
62 |
90 |
|
14.16–16.21 |
41 |
14 |
80 |
135 |
|
14.46–16.24 |
20 |
18 |
50 |
88 |
|
14.28–16.18 |
42 |
15 |
53 |
110 |
|
Total |
189 (29.35%) |
84 (13.04%) |
371 (57.61%) |
644 |
|
Average |
31.5 |
14 |
61.83 |
107.33 |
Table 4. Ceriagrion coromandelianum—duration and average timing (in
minutes) of the three stages of the final emergence recorded on site.
|
Time (h) |
Stage I |
Stage II |
Stage III |
Total |
|
12.43–15.08 |
65 |
10 |
70 |
145 |
|
12.56–15.06 |
47 |
11 |
72 |
130 |
|
13.06–15.47 |
46 |
13 |
102 |
161 |
|
14.10–16.30 |
35 |
25 |
80 |
140 |
|
14.23–16.38 |
47 |
16 |
72 |
135 |
|
Total |
240 (33.80%) |
75 (10.50%) |
396 (55.70%) |
711 |
|
Average |
48 |
15 |
79.2 |
142.5 |
For
figures & images – click here
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