Tiger beetles (Coleoptera: Cicindelidae) of ancient
reservoir ecosystems of Sri Lanka
Chandima Dangalle 1, Nirmalie
Pallewatta 2 & Alfried Vogler 3
1,2
Department of Zoology, Faculty of
Science, University of Colombo, Colombo 03, Sri Lanka
3
Department of Entomology, The Natural
History Museum, London SW7 5BD, United Kingdom
Email: 1 cddangalle@gmail.com (corresponding
author) 2 nirmalip@yahoo.com, 3 a.vogler@nhm.ac.uk
Date of publication (online): 26 April 2012
Date of publication (print): 26 April 2012
ISSN 0974-7907 (online) | 0974-7893 (print)
Editor: Anonimity
requested
Manuscript
details:
Ms # o2896
Received 29 July 2011
Final received 26 December 2011
Finally accepted 19 February 2012
Citation: Dangalle, C.,
N. Pallewatta & A. Vogler (2012). Tiger beetles (Coleoptera:Cicindelidae)
of ancient reservoir ecosystems of Sri Lanka. Journal of Threatened Taxa 4(4): 2490–2498.
Copyright: © Chandima
Dangalle, Nirmalie Pallewatta & Alfried Vogler 2012. Creative Commons
Attribution 3.0 Unported License. JoTT allows unrestricted use of this article
in any medium for non-profit purposes, reproduction and distribution by
providing adequate credit to the authors and the source of publication.
Author Details: Dr. Chandima Dangalle is a Senior Lecturer in Zoology,
attached to the University of Colombo, Sri Lanka. Her expertise lies in the
fields of Entomology and Molecular Biology. Her research focuses on collecting
baseline data on the distribution and habitat preferences of tiger beetles in
Sri Lanka. Further, she studies the evolution and phylogeny of the species
using mitochondrial DNA sequences of collected specimens. She conducted her PhD
in the Department of Zoology, University of Colombo, Sri Lanka and Department
of Entomology, Natural History Museum, London, United Kingdom.
Dr. Nirmalie Pallewatta is a Senior
Lecturer and the current head of the department of Zoology, University of
Colombo, Sri Lanka. A zoologist by training, she received her PhD in 1986 from
the Imperial College of Science, Technology and Medicine at the University of
London, United Kingdom.
Dr. Alfried Vogler works on the
molecular systematics of Coleoptera. He has a joint position at the
Natural History Museum and at Imperial College, London. Together with PhD
students and postdocs, he is currently studying basal relationships of
Scarabaeinae and Aphodiinae. He is also interested in the factors
determining the composition of dung beetle communities and the effect of
species interactions on the evolution of ecomorphological diversity.
Author Contribution: CD conducted
field studies in Sri Lanka and laboratory work in the Natural History Museum,
London, United Kingdom. She contributed towards research design and methodology
and writing of the paper. NP contributed towards formulating the initial
concept, research design and methodology and writing of the paper. AV
contributed by formulating the initial concept and research design.
Acknowledgments:
We wish to thank the National Science Foundation of Sri Lanka (Research
Grant No. RG/2003/ZOO/01) for funding the present study. We are also greatly
indebted to the Department of Zoology, University of Colombo; the Natural
History Museum of London, United Kingdom and the Department of Wildlife
Conservation of Sri Lanka. We are grateful to Prof. Nimal Dangalle, Department
of Geography, University of Kelaniya, Sri Lanka for his assistance in the preparation
of maps and locational lists.
Abstract: The
reservoir ecosystems of Sri Lanka are ancient man-made riparian habitats. Adequate food supply and suitable
climatic and soil parameters make these habitats ideal for tiger beetles. Twenty-six reservoirs were investigated
for the presence of tiger beetles, and four species were recorded: Calomera angulata
(Fabricius, 1798), Myriochila
(Monelica) fastidiosa (Dejean,
1825), Cylindera
(Oligoma) lacunosa (Putzeys, 1875) and Lophyra (Lophyra) catena
(Fabricius, 1775). Calomera angulata
is the most common species, occurring in the majority of reservoir
habitats. Key environmental
factors of climate and soil were examined and linked to habitat preferences of
tiger beetle species.
Keywords:
Coleoptera, Cicindelidae, habitat
preferences, reservoirs, tiger beetles.
For
figures, images, tables -- click here
INTRODUCTION
Tiger
beetles (Coleoptera: Cicindelidae) have been recorded from Sri Lanka since the
1860s. Identified species, their
distributions and habitats are given by Tennent (1860), Horn (1904), Fowler
(1912), Wiesner (1975), Naviaux (1984) and Acciavatti & Pearson
(1989). These records together
show 59 tiger beetle species from Sri Lanka, of which 39 are endemic.
The
majority of tiger beetles in Sri Lanka are terrestrial and diurnal, and are
included in the genera Cicindela,
Calochroa, Calomera, Lophyra, Jansenia, Cylindera, Myriochila, Hypaetha and
Callytron. These species occupy a variety of
habitats on the island such as riverine sandy areas, beaches and coastal areas,
lagoons by the ocean, forests, forest openings, wet rocks along water courses,
grasslands, fallow fields and road cuts (Wiesner 1975; Naviaux 1984; Acciavatti
& Pearson 1989). However, the
habitats of many species are unrecorded, and current localities of occurrence
are unknown.
Tiger
beetles are highly habitat-specific (Knisley & Hill 1992; Adis et al. 1998;
Morgan et al. 2000; Cardoso & Vogler 2005; Satoh et al. 2006; Pearson &
Cassola 2007). Human activities in
Sri Lanka have caused habitat loss, fragmentation and degradation, increasing
the risk of extinction for many species including endemics (IUCN 2006). Therefore, it is highly likely that
insect species with narrow habitat requirements, such as the tiger beetles,
will be threatened with extinction in the near future given the pressures of
development in the country. Therefore,
it is imperative that the current occurrence and status of tiger beetles be
investigated.
The
ancient man-made reservoir (tank) systems are a unique habitat in Sri Lanka,
dating back to about 2500 years
when the country had a hydraulic
civilization. They were built by the kings for irrigation purposes, domestic
and municipality needs, and flourish today in the ancient kingdoms of
Anuradhapura, Polonnaruwa and Sigiriya in the North-Central Province of the
country (Bandaragoda 2006). Over
30,000 reservoirs have been built in Sri Lanka, and some are still found
covered by thick jungle dotting the landscape all over the country, especially
in the dry zone (De Silva 1988). A
noteworthy, modern day feature of this unique system for water storage is that
it performs important roles in conservation of Sri Lankan biodiversity apart
from the original roles for which they were constructed. Although the reservoirs are man-made
they very often blend seamlessly with the natural environment and it is nearly
impossible to separate the man-made, reservoir-based agricultural environment
from the natural environment.
We
report here the first recorded occurrence of tiger beetle species from the
ancient reservoir ecosystems of Sri Lanka. The study reports the occurrence of four tiger beetle
species that have been previously found in other habitat types in Sri Lanka and
other countries, associated with reservoir habitats for the first time.
Further, we reveal the habitat preferences of the tiger beetle species associated
with the reservoir habitats of Sri Lanka.
METHODS AND MATERIALS
Study area
Twenty
six reservoirs were surveyed for the occurrence of tiger beetle species from
December, 2003 to November 2005.
Most of the reservoirs were located in the North-Central Province of the
country, while the other reservoirs were located in the North-Western,
Southern, Central and Western provinces of the island (Fig. 1, Table 1).
Measuring habitat variables of the
reservoirs
The
habitat variables of the climate and soil of the reservoirs in which tiger
beetles occurred
were measured as follows:
(i) Climate variables: The
ambient temperature, degree of solar radiation, relative humidity and wind
speed of the habitat were recorded using a portable integrated weather station
with optional sensors (Health EnviroMonitor, Davis Instrument Corp., Hayward,
CA, USA).
(ii) Soil variables: These
included the soil type/texture, using the sedimentation technique “soil
textural triangle” (Bierman 2007); soil colour, measured by comparison with a
Munsell soil colour chart; soil temperature, determined by using an Insert soil
thermometer (SG 680-10) ranging from -10 to 110 0C; soil pH,
determined by using a portable soil pH meter (Westminster, No. 259); soil
moisture, determined by selecting five random spots of a locality and
collecting samples down to a depth of 10cm and estimating the difference in
weight before and after oven drying to 107–120 0C in the
laboratory; and soil salinity, determined by a YSI model 30 hand-held salinity
meter.
Collection of beetles
Tiger
beetle species
were surveyed between 1000 –1500 hr at all localities. Adult tiger beetles were searched in
specific habitats including the bank of the reservoir, surrounding shrub area
near the reservoir, and off-road trails.
Beetles were collected using a standard insect net. Specimens were preserved in 96% ethanol
and stored at -200C after examining for morphological characters and
recording morphometric measurements.
Permission to enter various areas for tiger beetle collection and for
collecting specimens was obtained from the Department of Wildlife Conservation
of Sri Lanka.
Identification of tiger beetles
Taxonomic
keys of the Cicindela
of the Indian subcontinent by Acciavatti & Pearson (1989), descriptions of
Horn (1904) and Fowler (1912) were used to identify the species and
confirmation of identification was done through comparisons with specimens
available at the National Museum of Colombo and Natural History Museum (NHM),
London. Taxonomic names of
species, with the present nomenclatural changes, are based on Wiesner 1992,
except for the use of Calomera
instead of Lophyridia,
which is based on Lorenz (1998).
The
beetles were observed under a photomicrographic attachment which was also used
in photographing each specimen (Nikon AFX-DX, Tokyo, Japan).
RESULTS
Tiger
beetle species
were recorded from the sandy banks of eleven reservoirs of Sri Lanka. Most of the reservoirs were located in
the North-Central and North-Western provinces while a few were located in the
Central and Southern provinces (Image 1, Table 2).
Habitat variables of the reservoirs
Habitat
sampling from December 2003 to November 2005 revealed that tiger beetles occur
on sandy soils of reservoirs in areas of sparse vegetation. The beetles
exhibited a significant preference for sun-lit areas with high solar radiation
where climatic and soil temperatures were only slightly different. A soil moisture of 4.25±0.67 %
prevailed in the soils of the reservoirs which were more or less neutral with a
salinity of zero value. The
climatic and soil conditions of the reservoir habitats are given in Table 3.
Tiger beetle species recorded from the
reservoirs of Sri Lanka
Four
species of tiger beetles, Calomera
angulata (Fabricius, 1798), Myriochila (Monelica) fastidiosa (Dejean, 1825), Cylindera (Oligoma) lacunosa (Putzeys,
1875) and Lophyra
(Lophyra) catena (Fabricius,
1775) were recorded from the reservoir habitats of Sri Lanka (Table 2).
Calomera
angulata
(Fabricius, 1798) (Image 2)
Calomera angulata
was the most common tiger beetle species in reservoir ecosystems and dominated
all other species in terms of occurrence (Fig. 2). The species was found in nine out of 11 reservoirs and
formed large populations of a single species in five of the habitats
(Batalagoda Wewa, Kala Wewa, Mahakanadarawa Wewa, Parakrama Samudra, Thisa
Wewa), while in the other four tank systems (Devahuwa Wewa, Kandalama Wewa,
Nachchaduwa Wewa, Tabbowa Wewa) it co-occurred with either Myriochila (Monelica) fastidiosa, Cylindera (Oligoma) lacunosa or Lophyra (Lophyra) catena (Table
2). Even when co-occurring, Calomera angulata was
more abundant than the other species.
Myriochila
(Monelica) fastidiosa (Dejean,
1825) (Image 3)
Myriochila (Monelica) fastidiosa was
recorded from four reservoir ecosystems of Sri Lanka. In three reservoirs, Kandalama Wewa, Nachchaduwa Wewa,
Tabbowa Wewa, it co-occurred with Calomera
angulata, while a single population was found at Nuwara Wewa,
Anuradhapura (Table 2).
Cylindera
(Oligoma) lacunosa (Putzeys,
1875) (Image 4)
A
single specimen of Cylindera
(Oligoma) lacunosa was found
co-occurring with Calomera
angulata and Lophyra
(Lophyra)
catena at
Devahuwa Wewa, Central Province. C. lacunosa occupied
the wet sandy habitat most close to the water edge of the reservoir.
Lophyra
(Lophyra) catena (Fabricius,
1775)
(Image 5)
Lophyra (Lophyra) catena were
encountered at Devahuwa Wewa, Central Province and Ridiyagama Wewa, Southern
Province (Table 2). The species
occupied sandy bank area of the reservoir shaded by grasses and shrubs.
DISCUSSION
Reservoir
ecosystems of Sri Lanka are riparian habitats that were constructed by humans
about 2500 years ago (Bandaragoda 2006), and are integrated and inter-woven
with the natural environment. The
ecosystem consists of a reservoir, a sandy bank, a strip of trees downstream of
the reservoir that act as a wind breaking barrier, and paddy fields. The sandy bank formed along the margin
of the water level attracts many invertebrates due to accumulated organic
matter and high food supply. Such
riparian habitats are known to be preferred by tiger beetles not only because
of adequate food resources but also due to safety from predators and low human
disturbance (Bhargav & Uniyal 2008).
Tiger
beetles are known to be specialized species with narrow habitat requirements,
hence changes in the habitat can lead to their disappearance (Diogo et al.
1999). Morphological differences
between species of tiger beetles are apparently affected by selection for
specific habitat requirements (Cardoso & Vogler 2005). Therefore, the discovery of tiger
beetles in the reservoir ecosystems of Sri Lanka, identification of the species
and recordings of habitat conditions of the ecosystems is of utmost importance.
Tiger
beetles are predatory insects that prefer riverine habitats with sandy soils
and minimal vegetation, where periodic disturbance by wind and water removes
encroaching vegetation (Warren & Buttner 2008). Female tiger beetles are specific in choosing oviposition
sites, as larval stages are soil dwelling and spend their entire life in the
same location (Brust et al. 2006).
The larvae of Cicindela
hirticollis of Nebraska, USA are known to select burrow
locations with at least 7% soil moisture to avoid difficulties associated with
digging in loose, dry sand and to avoid abrasion of soft-bodied larvae.
Further, soil moisture is known to be necessary for cohesion of soil particles
and to prevent the collapse of the burrow walls (Brust et al. 2006). However, certain species that have
evolved as sand dune species are known to prefer a soil moisture of less than
4% and have higher amounts of cuticular hydrocarbons to avoid dessication
(Romey & Knisley 2002). The
tiger beetle species of the reservoir ecosystems of Sri Lanka were found on
soils of 4.25±0.67 % moisture, a value in between the above extremes. The colour of the soil on which they
occur is also known to be correlated with the structural colouration of
species, an apparent adaptation for remaining inconspicuous to natural enemies
reliant on visual cues (Seago et al.
2009). The
closely packed pits on the elytral surface of tiger beetles and the surface
microsculpture of the elytron is capable of reflecting wavelengths that create
a diffuse matte brown, green or similarly unsaturated hue; often matching the
colour of the surrounding soil (Seago et al. 2009).
The
soils of the reservoirs of Sri Lanka inhabited by cicindelids were mainly
brown, matching the bronze, copper-green and copper-brown colours of Calomera angulata,
Myriochila (Monelica) fastidiosa, Lophyra (Lophyra) catena and Cylindera (Oligoma) lacunosa. The
expanded white maculations on the elytra may have functioned in lowering the
body temperature making them able to forage longer without overheating.
According to Seago et al. (2009), Cicindela
formosa and Neocicindela
perhispida found on white beaches have expanded white
maculations that significantly lowers the body temperature and enabling them to
forage longer without overheating.
The environmental temperature of the reservoir ecosystems which was
34.27 ± 2.960C may also be suitable for the occurrence of tiger
beetles as ground temperature ranging from 32–330C is known to
be suitable for the activity and viability of tiger beetle populations, and a
temperature of 34 – 350C determined the greatest number of
matings in Cicindela
(Cephalota) circumdata leonschaeferi
Cassola (Eusebi et al. 1989).
Calomera angulata
was the most common tiger beetle species found in a majority of reservoir
ecosystems. When
co-occurring it was far more abundant than the other species and only single
specimens of L. catena
and C. lacunosa
were found co-occurring with C.
angulata at Devahuwa wewa. The low number of sympatric tiger beetle species in
reservoir habitats may be due to competition for food resources as all species
in these habitats had more or less similar mandible lengths (Table 4).
Calomera angulata, the key tiger
beetle species of the reservoir ecosystems of Sri Lanka, has been reported from
Sri Lanka as far back as 1904 (by Horn) and 1912 (by Fowler). However, it was identified as Cicindela sumatrensis
Herbst 1806 and was reported from riverine and coastal locations, and certain
other locations where the habitat is not defined. Wiesner (1975) and Acciavatti & Pearson (1989) have
recorded Calomera
angulata from India and Nepal, but have not recorded the
species from Sri Lanka. Wiesner
(1975) reports that the species can be found near the water’s edge on open,
moist sandy banks of rivers. More recently, Shook (1987) has reported Calomera angulata
from along river habitats in Thailand. However, Satoh & Hori (2004) define Calomera angulata
as a coastal tiger beetle occurring along the sea coast of Japan co-occurring
with other tiger beetle species in most instances.
Myriochila (Monelica) fastidiosa, a
species restricted to Sri Lanka and India has been cited in several works of
literature inhabiting grasslands, scrub forests, forest paths and old fields
(Horn 1904; Fowler 1912; Naviaux 1984; Acciavatti & Pearson 1989).
Lophyra (Lophyra) catena was first
recorded in Sri Lanka in 1860 by Tennent, and later by Horn (1904), Fowler
(1912), Naviaux (1984) and Acciavatti & Pearson (1989). Naviaux (1984) reported the species
from margins of rivers and lagoons by the ocean, as well as in large sunny
forest clearings. However, field
work done in the present study (December 2003 to November 2005) also revealed
the species from sandy lawns, foot paths and dry sand of coastal areas away
from the water.
Cylindera (Oligoma) lacunosa, a species
restricted to Sri Lanka and Tamil Nadu, India, has been reported from forest
openings of Sri Lanka (Horn 1904; Fowler 1912; Naviaux 1984; Acciavatti &
Pearson 1989).
None
of the above tiger beetle species are endemic to Sri Lanka and can be found in
the Indian subcontinent and countries of South East Asia. The vast majority of the endemic flora
and fauna of Sri Lanka are restricted to the wet zone of the country as are the
tiger beetle species (Dangalle et al. 2011a,b). Therefore, the reservoir habitats of Sri Lanka are perhaps
less important than other habitats for supporting endemic tiger beetle species. However, the reservoir habitats may
have facilitated the dispersion of tiger beetles within Sri Lanka and may have
played a role in facilitation of colonization of wet zone habitats by the
endemic tiger beetles, by provision of transitional habitats. The two endemic tiger beetle species, Cylindera (Ifasina) waterhousei and Cylindera (Ifasina) willeyi, reported
from the wet zone of Sri Lanka (Dangalle et al. 2011a,b) are species of the
genus Cylindera
which occurs in subtropical and temperate regions of Africa, Madagascar,
Eurasia, Asia and South-East Asia (Sota et al. 2011). According to Pearson & Ghorpade (1989) these taxa
dispersed to Sri Lanka using continuous forest habitats that were available
from south-eastern Asia to the Indian subcontinent, and traveled down the
Western and Eastern Ghats to reach central Sri Lanka which was continuous with
the southern division of the Western Ghats. The reservoir habitats may have provided a suitable habitat
for the dispersal of these small bodied beetles with weak flying
abilities. Similarly, as many
tanks are connected to a network of canals and streams and thus indirectly to
major rivers, it is tempting to theorize that coastal zone species would have
made the transition to reservoir habitats along such interconnected waterways.
More data on the occurrence and viability of tiger beetle populations along the
canal and stream networks of the ancient tank system needs to be collected.
In
conclusion our study reveals that reservoir (tank) habitats which are man-made
and dating back thousands of years have been colonized (‘invaded as new
habitat’) by tiger beetle species which are known to occupy other types of
habitats elsewhere in the world.
Suitable climatic and soil conditions of the locations have facilitated
the occurrence of tiger beetle species in these habitats and the study reports
the habitat preferences of the species of reservoir ecosystems of Sri Lanka.
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