Journal of Threatened Taxa |
www.threatenedtaxa.org | 26 November 2021 | 13(13): 20011–20018
ISSN 0974-7907 (Online) | ISSN 0974-7893
(Print)
https://doi.org/10.11609/jott.5458.13.13.20011-20018
#5458 | Received 06 October 2019 | Final
received 04 September 2021 | Finally accepted 22 September 2021
Diversity of aquatic insects and
biomonitoring of water quality in the upper Ganga River, a Ramsar
site: a preliminary assessment
Kritish De 1, Arkojyoti Sarkar 2, Kritika Singh 3,
Virendra Prasad Uniyal 4, Jeyaraj Antony Johnson 5 & Syed Ainul Hussain 6
1–6 Wildlife Institute of India, Chandrabani, Dehradun, Utarakhand
248001, India.
1 Present address: Department of
Life Sciences, Sri Sathya Sai University for Human Excellence, Navanihal, Okali Post, Kamalapur, Karnataka 585313, India.
1 kritish.de@gmail.com
(corresponding author), 2 arko.joti777@gmail.com, 3 kritika17singh@gmail.com,
4 uniyalvp@wii.gov.in, 5 jaj@wii.gov.in, 6
hussainsyedainul@wii.gov.in
Editor: Asheesh Shivam
Mishra, Nehru Gram Bharati (Deemed to be University), Prayagraj,
India. Date of publication: 26 November 2021
(online & print)
Citation: De, K., A. Sarkar, K. Singh,
V.P. Uniyal, J.A. Johnson & S.A. Hussain (2021). Diversity of aquatic insects and
biomonitoring of water quality in the upper Ganga River, a Ramsar
site: a preliminary assessment. Journal of Threatened Taxa 13(13): 20011–20018. https://doi.org/10.11609/jott.5458.13.13.20011-20018
Copyright: © De 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: This work was funded by the
National Mission for Clean Ganga, Ministry of Jal Shakti, Department of Water
Resources, River development and Ganga Rejuvenation, Government of India (Grant
No. B-02/2015-16/1259/NMCG-WIIPROPOSAL).
Competing interests: The authors
declare no competing interests.
Author details: Kritish De worked as project fellow at the Wildlife Institute of
India. Presently he is working as Assistant Professor at Sri Sathya Sai
University for Human Excellence. His research interests are biodiversity and
ecology. Arkojyoti Sarkar is working as project fellow.
His research interests are biodiversity and ecology. Kritika Singh worked as project intern. Her research
interest is monitoring of environmental health. Virendra
Prasad Uniyal is working as Scientist G. His research
interests are ecology and systematics of insects, bioindicators, biodiversity,
and ecological monitoring. Jeyaraj Antony Johnson is working as Scientist
E. His research interests are ecology and monitoring of aquatic ecosystems. Syed Ainul Hussain
worked as Scientist G. His research interests are aquatic ecology and
conservation biology.
Author contributions: KD—conceptualization, field work, formal analysis, writing original
draft; AS—field work, writing original draft; KS—field work, writing original
draft; VPU—supervision, review and editing the draft; JAJ—supervision, review
and editing the draft; SAH—supervision,
review and editing the draft, funding acquisition.
Acknowledgements: The authors are thankful to: the
National Mission for Clean Ganga, Ministry of Jal Shakti, Department of Water
Resources, River Development and Ganga Rejuvenation, Government of India for
sponsoring the work under the project ‘Biodiversity conservation and Ganga
Rejuvenation’; the director and dean, Wildlife Institute of India, for their
administrative support for the study; the Environment, Forest and Climate
Change Department, Government of Uttar Pradesh for necessary support during
fieldwork.
Abstract: Monitoring of freshwater
habitats through aquatic insects is widely used. A study was carried out in
March, 2019 at 14 sites in the Upper Ganga River between Brijghat
and Narora, a riverine Ramsar
site in India, to document the diversity of three major aquatic predatory
insect groups—Odonata, Coleoptera, and Hemiptera—and
determine their biomonitoring potential. The study recorded three species of Coleoptera, four Hemiptera, 14 dragonflies, and eight
damselflies. The Shannon diversity index (H′) ranged from 2.465 to 2.782, Pielou’s Evenness index (J′) from 0.841 to 0.894, and
Berger–Parker index of dominance (d) from 0.122 to 0.243. Families Libellulidae (Odonata), Coenagrionidae
(Odonata) and Gerridae (Hemiptera) had high relative
abundance and dominant status. The stream invertebrate grade number-average
level (SIGNAL2) score (for family) ranged from 2.316 to 3.174, lying within
quadrant 2 of the SIGNAL2 (family) quadrant diagram. This suggested that the
water in the area is likely to have high levels of turbidity, salinity, or
nutrients, caused naturally or by
anthropogenic activities, and the water has low levels of most toxic chemicals.
Keywords: Coleoptera, Hemiptera, Odonata, SIGNAL2
(family) score.
INTRODUCTION
Freshwater habitats occupy 1% of
the earth’s surface (Strayer & Dudgeon 2010), and in addition to supporting
many species freshwater ecosystems provide goods and services of critical
importance to human societies. Nevertheless, they are among the most heavily
altered ecosystems, with proportional loss of biodiversity (Geist 2011), owing
to human activities that have led to widespread habitat degradation, pollution,
flow regulation, water extraction, fisheries overexploitation, and alien
species introductions (Strayer & Dudgeon 2010). Alterations of natural flow
regimes by manmade dams, land use changes, river impoundments and water
abstraction often have profound impacts on lotic communities (Geist 2011).
Aquatic insects are an indispensable part of food webs and of nutrient cycling
in freshwater ecosystems, and they are essential components of the diets of
fish, amphibians and many birds and mammals (Morse 2017). Their abundance and
responses to changes in their environment make aquatic insects key indicators
for monitoring the effects of human activity on water quality (Adu & Oyeniyi 2019), and they
widely used for freshwater ecosystem monitoring (Souto
et al. 2019).
In India, 42 wetlands of
international importance (i.e., Ramsar sites) cover
1,081,438 ha according to the Ramsar Sites Information
Service (https://rsis.ramsar.org/sites/default
/files/rsiswp_search/exports/Ramsar-Sites-annotated-summary-India.pdf?1625598230).
Among these wetlands, information on aquatic insect communities and their
utility is scant. There are a few studies available on aquatic insect
communities of Indian Ramsar sites such as eastern
Kolkata wetlands in West Bengal (Sahaet al. 2007),
Pong Dam in Himachal Pradesh (Babu et al. 2009), Loktak Lake in Manipur (Takhelmayum
& Gupta 2011, 2015), Deepor beel
in Assam (Sharma & Sharma 2013; Choudhury & Gupta 2017), and Nalsarovar Bird Sanctuary in Gujarat (Rathod & Parasharya 2018).
The use of insects as
bioindicators is a low-cost strategy for preliminary assessments of the water
quality of inland freshwater bodies, as it avoids the use of expensive
analytical methods (Pal et al. 2012). The top predators among insects in
aquatic ecosystems include aquatic Coleoptera,
Hemiptera, and Odonata (Klecka & Boukal 2012). This study assessed diversity of these groups
in the upper Ganga River, a Ramsar site; the goal of
using them as indicators of water quality.
MATERIALS
AND METHODS
The study was conducted in an
85-km stretch of the river Ganga from Brijghat to Narora in Uttar Pradesh (Figure 1). This section of the
river was declared a Ramsar site in 2005 and is
generally characterized by shallow water, although some deep water pools are
present inhabited by conservative significant species such as Ganges River
Dolphin, Gharial, crocodiles, turtles, otters, 82 species of fish and more than
a hundred species of birds. The study was carried out during March 2019. The
study area was stratified into 14 sampling sites with a distance of ~5 km
between two sites and insect sampling was done at each site. At each study
site, sampling was done between 0930 h and 1130 h along the left bank (because
of accessibility to the river bank) of the main channel of the river Ganga.
To collect odonates,
a 100 m × 20 m transect (subdivided into 20 segments of 5 m) (Juen & De Marco 2011) was placed at each sampling site
parallel to and ~1 m beside the main river channel. Adult odonates
present in each of these segments were captured using insect collection nets
(mesh size 60 µm) and released after identification using published pictorial
field guides (Andrew et al. 2008; Subramanian 2009; Nair 2011). For Coleoptera and Hemiptera, a circular net (mesh size 60 μm) was dragged in the open water for one minute and continued three times per site (Subramanian
& Sivaramakrishnan 2007). All samples were
preserved in 70% ethanol and brought to the laboratory for further analysis.
They were later identified at species level using a stereo zoom microscope with
the help of taxonomic literature (Bal & Basu
1994a,b; Biswas & Mukhopdhyay 1995; Biswas et al.
1995; Chandra & Jehamalar 2012).
The aquatic insect data were
subjected to Shannon diversity index (H′), Pielou’s
evenness index (J′), and Berger–Parker index of dominance (d) index analysis.
The dominant status of the insects was calculated according to Engelmann’s scale
(1978) in which if relative abundance of a species is up to 1%, it is
considered as subrecedent; if between 1.1–3.1%, recedent; if between 3.2–10%, subdominant; if between
10.1–31.6 %, dominant, and if 31.7% or more then eudominant.
By evaluating comparative
performance of several aquatic health indices, Cox et al. (2019) found that the
stream invertebrate grade number-average level (SIGNAL2) is the most sensitive
index, family richness percentage is the most robust index, family richness and
family richness percentage are the best ranked indices for both measures of
usability; but Australian River Assessment System (AUSRIVAS OE50),
Ephemeroptera Plecoptera & Trichoptera
index (EPT), and Bray-Curtis index (BCI) have poor performance to asses river
health condition.
In this study, for the assessment
of the bioindicator potential of the insects, SIGNAL2 (family) score was used
which is a family-level water pollution index based on the known tolerances of
aquatic macro-invertebrate families to various pollutants which has a gradient
from 1 to 10 (ranging from a pollution tolerant to a pollution sensitive
community) (Chessman et al. 1995). The SIGNAL2 (family) scores were plotted in
a quadrant diagram (SIGNAL2 score in the y axis and the numbers of families in
the x axis) which includes four quadrants. The first quadrant indicates
favourable habitat and chemically dilute waters, the second quadrant indicates
high salinity or nutrient levels (may be natural), the third quadrant indicates
toxic pollution or harsh physical conditions and the fourth quadrant indicates
urban, industrial or agricultural pollution, or downstream effects of dams
(Chessman et al. 1995).
All the analyses were performed
in the software Past 3 (Hammer et al. 2001) and R 3.5.3 (R Core Team 2019).
RESULTS
A total of 29 species of aquatic
insects were recorded (Table 1), including three species of Coleoptera
belonging to two families, four species of Hemiptera belonging to four
families, and 22 species of Odonata belonging to three families. Among the odonates, 14 were dragonflies (Suborder Anisoptera)
and eight were damselflies (Suborder Zygoptera). Nine
species were recorded from all 14 sampling sites: Gerriss
pinolae Lethierry &
Severin, 1896; Anisops campbelli
Brooks, 1951; Brachythemis contaminata Fabricius, 1793; Diplacodes trivialis
Rambur, 1842; Orthetrum sabina
Drury, 1770; Trithemis aurora
Burmeister, 1839; Ceriagrion coromandelianum Fabricius,
1798; Pseudagrion decorum Rambur, 1842,
and Pseudagrion rubriceps
Selys, 1876.
The Shannon diversity index (H′)
ranged from 2.465 (at S8) to 2.782 (at S14) (mean= 2.579, SD= 0.086); Pielou’s evenness index (J′) was maximum at S7 (J’= 0.894),
and Berger-Parker index of dominance (d) ranged from 0.122 (S7) to 0.243 (S11)
(mean= 0.170, SD= 0.037). Variation of Shannon diversity index (H′), Pielou’s evenness index (J′), and Berger-Parker index of
dominance (d) are given in Figure 2.
For families, Gerridae
(Hemiptera) was dominant in >92 % of sampling sites, and Notonectidae
(Hemiptera) in >28 % of sites. Libellulidae
(Odonata) was eudominant in >64 % of sampling
sites and dominant in >35 % of sites, while Coenagrionidae
(Odonata) was eudominant in >71 % of the sampling
sites, and dominant in >28 % of sites. Dominance status in different sites
is given in Table 2.
The family richness and the family
richness percentage varies from 7 to 9 and 77.77 to 100 %, respectively.
Highest family richness and family richness percentage was found at S10.
The SIGNAL2 (family) score ranges
between 2.316 (S6) and 3.174 (S11) (mean= 2.579, SD= 0.086). The family richness,
family richness percentage and SIGNAL2 (family) score showed an increasing
trend in values from S1 to S14 (Figure 3).
The SIGNAL 2 quadrant diagram
plots SIGNAL 2 scores (on y axis) against numbers of aquatic invertebrate
families (on x axis). Each diagram has four quadrants which represent different
status of water and habitat qualities (Chessman 2003). In the present study,
the SIGNAL2 (family) score ranged from 2.316 to 3.174 (Figure 3) and fell
within the quadrant 2 (Figure 4).
DISCUSSION
Insects have the ability to move
from unfavourable habitats to favourable ones. If a habitat becomes polluted or
altered, tolerant species will thrive and sensitive ones will move to a more
suitable habitat (Medina et al. 2007). Thus habitat alternation, either by
natural process or by anthropogenic impacts, can shape invertebrate communities. Aquatic
macroinvertebrates constitute important components of their ecosystems, and
they exhibit differential tolerances to changes in environmental conditions (Adu & Oyeniyi 2019). In the
present study, three species of Coleoptera from two
families, four species of Hemiptera from four families, and 22 species of
Odonata from three families were recorded. The coleopterans included predaceous
diving beetles (family Dytiscidae) and water
scavenger beetles (family Hydrophilidae). The
hemipteran group included water bugs (family Belostomatidae),
water striders (family Gerridae), water scorpions
(family Nepidae) and backswimmers (family Notonectidae), and the odonates
included dragonflies and damselflies.
In the present study Shannon
diversity index (H′), Pielou’s Evenness index (J′)
and Berger-Parker index of dominance (d) did not differ much between study
sites, probably because of uniform geomorphological features of the area, as
geomorphological heterogeneity plays a major role in determining species
richness (Nichols et al. 1998). Libellulidae, Coenagrionidae, and Gerridae had
high relative abundance and dominant status, probably because of their ability
to tolerate a wide range of environmental factors (Spence 1983; Chang et al.
2014).
The SIGNAL 2 result suggested
that the water of the study area was likely to have higher levels of turbidity,
salinity or nutrients, which was perhaps caused either naturally, because of local
geology and soil types, or as a result of human activities and physical
conditions. Toxic chemicals were not present in large amounts (Chessman 2003).
The family richness, family
richness percentage and SIGNAL2 (family) score showed an increase in values
towards the Narora barrage, probably because of the
increase in water quantity (as the barrage stores more water) which directly
affects the physiochemical properties of the water and habitat structures.
The upper Ganga Ramsar site is facing stress from anthropogenic pressure (Kuniyal 2013; Pandey & Sharma 2013). The study stretch
between the Brijghat and Narora,
the Ganga is characterized by the presence of agricultural lands and numerous ghats (steps leading down to the river) with religious and
tourism importance on both the banks. Local people use the river bank and water
for bathing, cremation and other religious activities. Activities like cattle
grazing and fishing occur throughout the year. As a result, the river is
exposed to various threats like waste discharge, sewage disposal, agricultural
runoff, fishing, and river bank erosion.
While there is some regional
information, knowledge remains limited concerning the natural ranges and
ecology of species found in the Ganga (Nautiyal et
al. 2014). Long-term seasonal monitoring of the physiochemical properties of
the water, coupled with assessment of faunal and floral diversity as well as
socio-economic factors influencing the conditions of the area is recommended in
order to arrive at better management strategies.
Table 1. List of Coleoptera,
Hemiptera, and Odonata recorded from across the study area in different sites
of Upper Ganga Ramsar site (+ represents presence and
- represent absence).
Species |
Sampling sites |
|||||||||||||
|
S1 |
S2 |
S3 |
S4 |
S5 |
S6 |
S7 |
S8 |
S9 |
S10 |
S11 |
S12 |
S13 |
S14 |
Order: Coleoptera |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
Family: Dytiscidae (Predaceous Diving Beetle) |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
1. Cybister
limbatus (Fabricius,
1775) |
+ |
- |
- |
- |
+ |
+ |
+ |
+ |
- |
+ |
+ |
+ |
+ |
- |
2. Eretes
sticticus (Linnaeus, 1767) |
- |
+ |
- |
- |
+ |
- |
+ |
- |
+ |
+ |
+ |
+ |
- |
- |
Family: Hydrophilidae
(Water Scavenger Beetle) |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
1. Hydrophilus
senegalensis (Percheron 1835) |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
- |
+ |
+ |
+ |
Order: Hemiptera |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
Family: Belostomatidae (Water Bug) |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
1. Diplonychus
rusticus (Fabricius,
1781) |
- |
+ |
+ |
+ |
- |
+ |
- |
+ |
- |
+ |
+ |
+ |
- |
- |
Family: Gerridae (Water
Striders) |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
1. Gerris
spinolae Lethierry
& Severin, 1896 |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
Family: Nepidae (Water
Scorpion) |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
1. Ranatra
elongate Fabricius, 1790 |
+ |
- |
+ |
+ |
+ |
+ |
+ |
- |
+ |
+ |
+ |
- |
+ |
+ |
Family: Notonectidae
(Backswimmers) |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
1. Anisops
campbelli Brooks, 1951 |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
Order: Odonata |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
Suborder: Anisoptera
(Dragonflies) |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
Family: Gomphidae |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
1. Platygomphus
dolabratus Selys,
1854 |
- |
- |
- |
- |
- |
- |
- |
- |
- |
- |
+ |
- |
- |
+ |
2. Ictinogomphus
rapax Rambur, 1842 (Indian Common Clubtail) |
- |
- |
- |
- |
- |
- |
- |
- |
- |
+ |
+ |
+ |
+ |
+ |
Family: Libellulidae
Leach, 1815 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
1. Acisoma
panorpoides Rambur, 1842 (Trumpet Tail) |
+ |
+ |
+ |
- |
+ |
+ |
+ |
+ |
+ |
- |
+ |
+ |
+ |
+ |
2. Brachydiplax
sobrina Rambur, 1842 (Little Blue Marsh Hawk) |
- |
- |
- |
- |
- |
- |
- |
- |
- |
- |
- |
+ |
- |
- |
3. Brachythemis
contaminate Fabricius, 1793 (Ditch Jewel) |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
4. Crocothemis
servilia Drury, 1770 (Ruddy Marsh Skimmer) |
+ |
- |
- |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
- |
+ |
+ |
5. Diplacodes
trivialis Rambur, 1842 (Blue Ground Skimmer) |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
6. Neurothemis
tullia (Drury, 1773) (Pied Paddy Skimmer) |
- |
- |
- |
- |
- |
- |
- |
- |
- |
- |
- |
- |
- |
+ |
7. Orthetrum
sabina Drury, 1770 (Green Marsh Hawk) |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
8. Pantala
flavescens Fabricius,
1798 (Wandering Glider) |
+ |
- |
+ |
+ |
+ |
+ |
+ |
- |
+ |
+ |
- |
- |
- |
+ |
9. Rhyothemis
variegate Linnaeus, 1763 (Common Picturewing) |
- |
- |
- |
- |
+ |
- |
- |
- |
+ |
- |
- |
+ |
+ |
+ |
10. Tramea
basilaris Palisot de Beauvois, 1805 (Red Marsh Trotter) |
- |
+ |
+ |
+ |
+ |
- |
- |
+ |
- |
- |
+ |
- |
+ |
+ |
11. Trithemis
aurora Burmeister, 1839 (Crimson Marsh Glider) |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
12. Urothemis
signata Rambur, 1842 (Greater Crimson Glider) |
- |
+ |
- |
+ |
+ |
+ |
+ |
+ |
+ |
- |
- |
- |
+ |
+ |
Suborder: Zygoptera
(Damselflies) |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
Family: Coenagrionidae |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
1. Agriocnemis
lacteolaSelys,
1877 (Milky Dartlet) |
+ |
+ |
- |
- |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
2. Agriocnemis
pygmaea Rambur, 1842 (Pygmy Dartlet) |
+ |
- |
+ |
- |
- |
+ |
- |
+ |
+ |
+ |
+ |
- |
- |
+ |
3. Amphiallagma
parvumSelys,
1876 (Azure Dartlet) |
+ |
+ |
+ |
+ |
+ |
- |
+ |
- |
- |
- |
- |
- |
+ |
+ |
4. Ceriagrion
coromandelianum Fabricius,
1798 (Coromandel Marsh Dart) |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
5. Ischnura
nursei Morton, 1907 (Pixie Dartlet) |
- |
- |
- |
+ |
- |
+ |
- |
- |
+ |
+ |
- |
- |
- |
+ |
6. Ischnura
rubilioSelys,
1876 (Western Golden Dartlet) |
+ |
+ |
+ |
+ |
- |
+ |
+ |
- |
+ |
+ |
+ |
+ |
- |
+ |
7. Pseudagrion
decorum Rambur, 1842 (Three Lined Dart) |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
8. Pseudagrion
rubriceps Selys, 1876
(Saffron Faced Blue Dart) |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
+ |
Table 2. Relative abundance (RA) and dominance status
(DS) of different families of aquatic insects in different sampling sites (S1
to S14) of the upper Ganga River, a Ramsar site
according to Engelmann’s scale (1978)
|
|
Dytiscidae |
Hydrophilidae |
Belostomatidae |
Gerridae |
Nepidae |
Notonectidae |
Gomphidae |
Libellulidae |
Coenagrionidae |
S1 |
RA |
1.515 |
1.515 |
|
21.212 |
1.515 |
9.848 |
|
34.091 |
30.303 |
DS |
Recedent |
Recedent |
|
Dominant |
Recedent |
Subdominant |
|
Eudominant |
Dominant |
|
S2 |
RA |
1.724 |
1.724 |
1.724 |
12.069 |
|
0.862 |
|
44.828 |
37.069 |
DS |
Recedent |
Recedent |
Recedent |
Dominant |
|
Subrecedent |
|
Eudominant |
Eudominant |
|
S3 |
RA |
|
2.069 |
1.379 |
18.621 |
0.69 |
12.414 |
|
30.345 |
34.483 |
DS |
|
Recedent |
Recedent |
Dominant |
Subrecedent |
Dominant |
|
Dominant |
Eudominant |
|
S4 |
RA |
|
0.758 |
1.515 |
19.697 |
3.03 |
9.091 |
|
30.303 |
35.606 |
DS |
|
Subrecedent |
Recedent |
Dominant |
Recedent |
Subdominant |
|
Dominant |
Eudominant |
|
S5 |
RA |
2.564 |
1.709 |
|
16.239 |
0.855 |
7.692 |
|
45.299 |
25.641 |
DS |
Recedent |
Recedent |
|
Dominant |
Subrecedent |
Subdominant |
|
Eudominant |
Dominant |
|
S6 |
RA |
0.735 |
2.206 |
0.735 |
15.441 |
0.735 |
12.5 |
|
33.088 |
34.559 |
DS |
Subrecedent |
Recedent |
Subrecedent |
Dominant |
Subrecedent |
Dominant |
|
Eudominant |
Eudominant |
|
S7 |
RA |
3.053 |
1.527 |
|
11.45 |
2.29 |
7.634 |
|
39.695 |
34.351 |
DS |
Recedent |
Recedent |
|
Dominant |
Recedent |
Subdominant |
|
Eudominant |
Eudominant |
|
S8 |
RA |
1.923 |
1.923 |
0.962 |
14.423 |
|
0.962 |
|
40.385 |
39.423 |
DS |
Recedent |
Recedent |
Subrecedent |
Dominant |
|
Subrecedent |
|
Eudominant |
Eudominant |
|
S9 |
RA |
0.73 |
1.46 |
|
15.328 |
2.19 |
12.409 |
|
43.066 |
24.818 |
DS |
Subrecedent |
Recedent |
|
|
Recedent |
Dominant |
|
Eudominant |
Dominant |
|
S10 |
RA |
3.008 |
0.752 |
0.752 |
16.541 |
2.256 |
10.526 |
0.752 |
27.82 |
37.594 |
DS |
Recedent |
Subrecedent |
Subrecedent |
Dominant |
Recedent |
Dominant |
Subrecedent |
Dominant |
Eudominant |
|
S11 |
RA |
1.429 |
|
1.429 |
24.286 |
1.429 |
2.857 |
5 |
30 |
33.571 |
DS |
Recedent |
|
Recedent |
Dominant |
Recedent |
Recedent |
Subdominant |
Dominant |
Eudominant |
|
S12 |
RA |
2.308 |
1.538 |
1.538 |
23.846 |
0 |
9.231 |
1.538 |
35.385 |
24.615 |
DS |
Recedent |
Recedent |
Recedent |
Dominant |
|
Subdominant |
Recedent |
Eudominant |
Dominant |
|
S13 |
RA |
1.835 |
2.752 |
|
14.679 |
2.752 |
0.917 |
0.917 |
39.45 |
36.697 |
DS |
Recedent |
Recedent |
|
Dominant |
Recedent |
Subrecedent |
Subrecedent |
Eudominant |
Eudominant |
|
S14 |
RA |
|
1.527 |
|
13.74 |
3.053 |
9.924 |
3.053 |
29.771 |
38.931 |
DS |
|
Recedent |
|
Dominant |
Recedent |
Subdominant |
Recedent |
Dominant |
Eudominant |
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