The current occurrence,
habitat and historical change in the distribution range of an endemic tiger
beetle species Cicindela(Ifasina) willeyi Horn
(Coleoptera: Cicindelidae) of Sri Lanka
Chandima
Dangalle 1, Nirmalie Pallewatta 2 & Alfried Vogler3
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@yahoo.com (corresponding author), 2nirmalip@yahoo.com, 3 a.vogler@nhm.ac.uk
Date of publication (online): 26 February 2011
Date of publication (print): 26 February 2011
ISSN 0974-7907 (online) | 0974-7893 (print)
Editor: K.A. Subramanian
Manuscript
details:
Ms
# o2501
Received
02 July 2010
Final
received 29 December 2010
Finally
accepted 05 January 2011
Citation: Dangalle,
C., N. Pallewatta & A. Vogler (2011). The current occurrence, habitat and historical change in
the distribution range of an endemic tiger beetle species Cicindela (Ifasina) willeyi Horn (Coleoptera:
Cicindelidae) of Sri Lanka. Journal of Threatened Taxa 3(2): 1493-1505.
Copyright: © Chandima
Dangalle, Nirmalie Pallewatta & Alfried Vogler 2011. 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. 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 and in evolution and
phylogeny of the species. Dr.
Dangalle 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 Dr. Pallewatta
received her Ph.D. in 1986 from the Imperial College of Science, Technology and
Medicine at the University of London, U.K.
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,
U.K. 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, U.K. 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 molecular sequencing methodology and analyses.
Acknowledgements: We wish to thank the
Department of Wildlife Conservation, Ministry of Environment, Sri Lanka for
providing permits necessary for field work and export of specimens. 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. This work has been supported by grant number RG/2003/ZOO/01
from the National Science Foundation of Sri Lanka.
Abstract: The current occurrence,
habitat and historical change in distributional range are studied for an
endemic tiger beetle species, Cicindela (Ifasina) willeyi Horn of Sri Lanka. At present, the species is only
recorded from Maha Oya (Dehi Owita) and Handapangoda, and is absent from the
locations where it previously occurred. The current habitat of the species is explained using
abiotic environmental factors of the climate and soil recorded using standard
methods. Morphology of the species
is described by studying specimens using identification keys for the genus and
comparing with specimens available at the National Museum of Colombo, Sri
Lanka. The DNA barcode of the
species is elucidated using the mitochondrial CO1 gene sequence of eight
specimens of Cicindela
(Ifasina) willeyi. The study
suggests that Maha Oya (Dehi Owita) and Handapangoda are suitable
habitats. However, its presence in
only two locations and its absence from locations where it previously occurred
highlights the need for conserving the natural habitats at Maha Oya (Dehi
Owita) and Handapangoda, and the necessity of further studies of this kind.
Keywords: Conservation, DNA barcode, habitat preferences, taxonomy, tiger
beetles.
For figures, images, tables --
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INTRODUCTION
Tiger Beetles (Coleoptera:
Cicindelidae) are a group of attractive, fast-flying and fast-running insect
predators that occur in many diverse habitats around the world (Pearson &
Cassola 2007). The ease with which
most species can be found and identified in the field, their habitat
specificity and their value as indicators of habitat health and biodiversity
have generated considerable interest among amateurs and professional biologists
alike (Pearson & Cassola 2007). As a result, a total of 2,559 species have been described world wide
since the first eight cicindelid species were identified by Carl Von Linné in
the 18th century (Pearson & Cassola 2005). Most tiger beetle species (29% of
described species) are found in the Oriental (Indo-Malaysian) region of the
world, while northeastern India has the highest recorded number of species in a small area which
is an 80km stretch between Siliguri and Darjeeling of West Bengal (Pearson
1988; Pearson & Cassola 2005). Sri Lanka is also attributed with a high number of cicindelid species
and ranks amongst the top 30 countries of the world with the highest number of
species (Cassola & Pearson 2000).
The existing literature
(Tennent 1860; Horn 1904; Fowler 1912; Naviaux 1984; Acciavatti & Pearson
1989), and collections available at the National Museum of Colombo and Natural
History Museum of London, documents 54 species of tiger beetles from Sri Lanka
belonging to five genera (Table 1), Cicindela, Tricondyla, Derocrania, Collyris andNeocollyris. The majority of species are included in
the genus Cicindelawhich consists of 13 subgenera, Ancylia, Calochroa, Lophyra, Lophyridia, Jansenia,
Oligoma, Cosmodela, Eugrapha, Monelica, Myriochile, Hypaetha, Callytron andIfasina(Table 1). Subgenus Ifasina is
attributed with the highest number of species (7) out of which five species are
endemic to the island.
We report in this paper the
first recorded occurrence of an endemic tiger beetle species, Cicindela
(Ifasina) willeyi, from two locations of Sri Lanka and the habitat
preferences of the species.
METHODS AND MATERIALS
Tiger beetles were surveyed
in 94 localities of Sri Lanka from May 2002 to December 2005 (Fig. 1, Table
2). The localities for collecting
tiger beetles were selected using information based on previous publications of
cicindelid species of Sri Lanka and information based on the different habitat
types of the family on a global scale.
Field studies were conducted
at Maha Oya (Dehi Owita) in August 2003 and 2004, and at Handapangoda in July
2003 and 2004.
Study Area: Maha Oya is a stream located
at Dehi Owita (6057’91”N & 80016’44”E, elevation
6.7m) in the Kegalle District, Sabaragamuwa Province of Sri Lanka. It covers an area of 17km2and encompasses the Grama Niladhari divisions of Dehi Owita, Debagama and
Thimbiripola. The stream is a
branch of the much larger Kelani River which starts from the Sri Pada Mountain
range and flows in to the ocean at Colombo. The water of Maha Oya is slow flowing and runs parallel to a
large sandy bank with 167m elevation (Fig. 2 & Image 1).
The stream at Handapangoda (6047’05N
& 80008’03E; elevation 23m) is located near a rubber estate in
the Kalutara District, Western Province of Sri Lanka. It covers an area of 11km2 and encompasses the
Grama Niladhari divisions of Handapangoda, Arakawila, Menerigama and
Kurana. It is a branch of Kalu
Ganga, which is the second largest river of Sri Lanka and is restricted
entirely to the wet zone of the country. The water of the stream is slow flowing and runs parallel to a narrow
sandy bank that consists mostly of rocks (Fig. 2 & Image 2).
Collection of Beetles: Four visits were made to each
site for surveying adult tiger beetles. Three belts of land of 100 to 150 m were selected on the sandy bank of
Maha Oya,while two belts were selected on the bank of Handapangoda. All sites were
sampled over five hours each day between 1000 to 1400 hr.
Beetles were searched for in
specific habitats and their immediate surroundings e.g. shrub area near the
sandy sections of the stream, rocky substrata, pathways and trails. When encountered tiger beetles were
rapidly counted at a given site using a hand tally counter (Wagtech, UK) and a sample was collected using a
standard insect net. The number of beetles and the sex ratio of those that
could be observed under field conditions and from collected specimens were
recorded.
Specimens were collected and
preserved in 96% ethanol and stored at -200C for subsequent
identification.
Measuring Habitat Variables: The following habitat
variables of the study sites were recorded.
(i) Weather variables: The
ambient temperature, degree of solar radiation, relative humidity and wind
speed of the habitat were recorded using a portable integrated weather station
(Health EnviroMonitor, Davis Instrument Corp., Hayward, CA, USA) with optional
sensors.
(ii) Soil characteristics: These
included the soil group (determined by using the generalized soil map of Sri
Lanka by Moorman & Panabokke 1961); soil type / texture (determined by the
sedimentation technique using the “soil textural triangle” as the reference
(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 to 120 0C in the laboratory.); soil
salinity (determined by a YSI model 30 hand-held salinity meter).
Determination of
morphological characteristics of tiger beetles: Morphological characters important in identification and for the
purpose of establishing characteristics of this species were also noted as
follows.
(i) Body weight: Each beetle
was weighed to the nearest mg on an analytical balance (Chyo JL180, Chyo
Balance Corp., Japan).
(ii) Body length: Estimated
by measuring the length from the frons of the head to the elytral apex when the
head was in the normal feeding position. The spines on the caudal end of the
elytra were disregarded. Measurements were taken using a dissecting microscope (Nikon Corporation
SE., Japan) with the aid of an eyepiece graticule (Nikon, Tokyo, Japan)
calibrated by an objective micrometer (Olympus, Japan).
Body lengths of beetles were
categorized as follows, based on size classes relevant for this group of
insects (Acciavatti & Pearson 1989).
1. very small – less
than 8mm
2. small – 8 to 10 mm
3. medium – 10 to 15 mm
4. large – 15 to 20 mm
5. very large – more
than 20mm
(iii) Mandible length /
chord: The distance from the articulation point of the left mandible to the tip
were measured under a dissecting microscope (Nikon Corporation SE, Japan) with
the aid of an eyepiece graticule (Nikon, Tokyo, Japan) calibrated by an
objective micrometer (Olympus, Japan). Only specimens with undamaged or not
noticeably worn mandibles were used (Pearson & Juliano 1993; Satoh &
Hori 2004).
(iv) Other characters: In
addition to the above characters the following features were also recorded for
each specimen.
1. Colouration – Dorsal and ventral colouration of the
body and its metallic or iridescent appearance was noted as was the colour of
the elytral maculae, pits, eyes, antennae, mandibles, labrum and legs.
2. Number of rugae between
eye and vertex.
3. Labrum – The shape
of the labrum and the number of labral setae.
4. Distribution of setae on
body.
5. Distribution of pits on
body.
The above characters were
observed under a photomicrographic attachment (Microflex AFX – DX, Nikon
Corporation, Tokyo, Japan) which was also used in photographing each specimen
(dorsal view, ventral view, lateral view, other important features). Additional photographs of species were
also taken by using the computer software programme Auto Montage (facilitated
with a SMC Pentax – FA macro camera) available at the entomology
laboratory of the Natural History Museum, London.
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.
DNA Sequencing: DNA analysis was carried out
to determine the DNA barcode of the species. Eight specimens from the two known
populations (Maha Oya and
Handapangoda) were used for DNA extraction and sequencing. DNA was extracted
from the abdominal region of the beetle using the DNeasy protocol (July
2003). Voucher specimens were
deposited in the Entomology Collection of the Department of Zoology, University
of Colombo.
A ~810 bp region of the 3’
end of the cytochrome oxidase 1 (CO1) gene was amplified using primers M202
(forward, 5’ - CAA CAT TTA TTT TGA TTT TTT GG - 3’, alias Jerry; Simon et al. 1994) and
M70 (reverse, 5’ - TCC ATT GCA CTA ATC TGC CAT ATT A - 3’, alias Pat; Simon et
al. 1994).
Standard PCR amplifications
included 5µl of NH4 buffer, 0.5µl of each dNTP, 1µl of each primer,
0.1µl of TAQ polymerase and 2.5µl of MgCl2 in a 50µl reaction
volume. PCR amplifications were
carried out on a GeneAmp PCR System 9700 thermal cycler (Applied Biosystems,
California, USA) for one cycle of (940C, 2.5 mins.; 470C,
0.5 mins.; 72ºC, 11 mins.; 40C, α) for 40 cycles. The PCR
products were purified using the UltraClean PCR clean-up DNA purification kit
(MoBio Laboratories Inc., Carlsbad, CA, USA) according to the manufacturer’s
protocol.
Sequencing was performed for
10µl of cleaned PCR product using the ABI Prism Big Dye Terminator Cycle
sequencing kit (PE Applied Biosystems, Foster City, CA, USA). PCR primers were used as sequencing
primers and each fragment was sequenced on both strands. The reaction products
were purified by ethanol precipitation and sequenced on ABI 373 (version 3.0)
automated DNA sequencer. Sequence files were edited using SEQUED version 1.0.3
(Applied Biosystems) and a consensus of bidirectional sequencing was
determined.
RESULTS
Cicindela (Ifasina) willeyi H. was the only tiger beetle
species observed at Maha Oya (Dehi Owita) and Handapangoda. Fifty beetles of Cicindela (Ifasina) willeyi. were observed
from Maha Oya (Dehi Owita) and five beetles were collected for morphometric and
morphological characterization. Ten specimens of the species were recorded from Handapangoda and three
were collected for characterization.
Taxonomy of Cicindela
(Ifasina) willeyi Horn, 1904: The description given in Horn (1904), Fowler (1912) and Acciavatti
& Pearson (1989) and the
specimens available at the National Museum, Colombo matches the features
observed in the specimens collected from Maha Oya (Dehi Owita) and Handapangoda
sites.
Diagnosis: Cicindela (Ifasina) willeyi H. closely resembles the
other allied species C. (I.) waterhousei Horn, C. (I.) dormeri Horn and C. (I.) ganglbaueri Horn that are also
endemic to Sri Lanka. However, the
species can be distinguished by the presence of an elytral humeral dot and
three spots that lie medially in a line on elytra; more excavate forehead
between the eyes; narrow, conical pronotum with a dorsal bulge that is
undivided by a short medial line (Image 3).
Description: Body small with an average
length of 9.19 ± 0.56 mm and an
average body weight of 27.45 ± 1.94 mg (n = 8) (Table 3). Dorsal region
of head, pronotum and base of eyes shiny copper green in colour. Elytra metallic brown with a humeral
dot, and three spots medially in a line. Spots yellowish-white in colour. First and third spot circular and small. Middle spot large, ovate or pear-shaped. Ventral side of body iridescent
bluish-green with white setae. Labrum short, broad and brown in colour with
eight or nine submarginal setae. Mandibles dark brown with left mandible having
an average length of 1.90 ± 0.54 mm (n = 3). Pronotum with a
medial dorsal bulge that is undivided by a medial line. Legs shiny blue-green in colour (Image
3).
Habitat of C. (I.) willeyi:Beetles were
found on a sandy bank at Maha Oya (Dehi Owita) which is a shallow stream found
in the Sabaragamuwa Province. The
periphery of the sandy bank consisted of shrub type vegetation of Gahala Colocasia
esculenta, Bamboo Bambusa vulgaris and long grasses that provided shade to the
habitat. Climatic and soil conditions of the site are given in Tables 4 and 5.
The climatic and soil
conditions at Handapangoda were more or less similar to that of Maha Oya (Dehi
Owita) (Tables 4 & 5). However, the beetles at Handapangoda were mainly
found on the moist rocks on the sandy bank. Further, the habitat at Handapangoda was more better shaded
than that at Maha Oya due to the large trees found on the bank.
Historical change in
distribution of C.
(I.) willeyi: The species has been reported from the Central Province of the
island and Labugama, Western Province from as far back as 1904 (Horn 1904;
Fowler 1912; Acciavatti & Pearson 1989; collection of the National Museum
of Colombo, Sri Lanka). However,
it has not been studied thereafter, and although it’s biology is well known,
data on present distribution and habitat is lacking. According to the results of the current study, at present it
occupies two habitats in Maha Oya (Dehi Owita) area and Handapangoda area. Investigations in the present study
have further revealed that the species is absent from its former locations of
Central Province and Labugama which have a geographical area of about 5583.5km2and is now restricted to an area of about 28km2 in Maha Oya (Dehi
Owita) and Handapangoda (Table 6).
Therefore, a loss of its
present habitats where it is restricted in distribution may threaten the
survival of the species and qualify it for a threatened category in the near
future.
DISCUSSION
Cicindela (Ifasina) willeyi is an endemic tiger beetle
species of Sri Lanka that was first described by Horn (1904) from the Central
province of Sri Lanka, which was subsequently confirmed by Fowler (1912) and
later by Acciavatti & Pearson (1989). The National Museum of Colombo houses a specimen of the species
collected from Labugama in the Western Province of the country. However, the present distribution of
this endemic species has been uncertain and habitat characteristics and
preferences were completely unknown. The present study reveals the species from Maha Oya (Dehi Owita)
(Sabaragamuwa Province), and Handapangoda (Western Province), localities from
which it had not been previously recorded. Further, extensive field work
carried out in riparian habitats of Meewathura, Kandy; Ma Oya, Mawanella; Pundalu Oya, Kotmale Oya, Ramboda and Silverfalls,
Nuwara Eliya (Central Province); and Labugama (Western Province) during the
study period confirms the absence of the species from its previous localities
of occurrence (Fig. 1). This knowledge on the past and present distribution of C. (I.) willeyiindicates the possibility of a historical change in the distributional range of
the species.
Various factors including
urbanization, increased intensity of recreational use of beaches, increased
off-road vehicle traffic, conversion of the habitat to a dumping ground for
automobile parts and construction material, increased vegetation encroachment
that eliminates open areas and inundation of habitat caused by the construction
of dams have been used to explain historical range changes in tiger beetles in
the United States (Knisley & Hill 1992; Kritsky et al. 1996; Knisley &
Fenster 2005; Pearson et al. 2006; Simmons 2007).
Therefore, it is possible
that C. (I.)
willeyi extirpated from the localities in the Central Province and
Labugama, and inhabited the area in Handapangoda and Maha Oya (Dehi
Owita). Even though, the reason
for this is not precisely known, possible events of the past can be suggested.
The Mahaweli Development
Programme was initiated in Sri Lanka in 1979 to fulfill the water requirements
for agriculture, industrial and domestic use, develop hydropower and reduce
flood peaks. As a result of this
programme, four major reservoirs and dams, namely, Kotmale, Victoria, Randenigala
and Rantambe were built along the
main stem of the river that lies in the Central Province (Dayawansa 2008).
Building of a dam is known to reduce and corrode soil and rock along the river
as massive deposits of soil are left within the reservoir. The massive cobblestones, sand and
crushed stones held back by the dam and the transformation of the once tortuous
riverbank and riverway to a relatively straight river course results in the
loss of habitat for many invertebrates who survive in this environment (Mao
& Zheng 2006). During reservoir filling the river and any associated
wetland areas become inundated. Riffles, runs and pools of the river are lost
beneath the rising waters, leading to the extirpation (or extinction) of
habitat sensitive riverine species with tightly defined niche requirements
(McAllister et al. 2001). Likewise, reservoir and dam construction along the Mahaweli River has
resulted a considerable impact on river morphology and has inundated a vast
area of land in the Central Province of the island. Aerial photographs obtained in 1985 and 2003, and
topographical maps of the Survey Department of Sri Lanka, present encroachments
towards the river across its floodplain area in Gohagoda, Central
Province. Further, riparian areas
in Peradeniya (Central Province) have changed to residential urban areas with
considerable impacts of waste disposal (Dayawansa 2008).
It is also recorded that gold
has been mined in the past from a concordant quartz reef in Central Sri Lanka
(Nawaratne & Dissanayake 2001), and several gemming grounds are located in
Nuwara Eliya, Horton Plains, Hatton and Kandy (Herath 1984).
Therefore, it is strongly
possible that C.
(I.) willeyi went locally
extinct from its historic sites in the Central Province of Sri Lanka and
invaded the riparian habitats of Maha Oya (Dehi Owita) (Sabaragamuwa Province) and
Handapangoda (Western Province) due to the unsuitability and loss of its former
habitats. The presence of
populations of the species at Maha Oya and Handapangoda, indicate the
suitability of the habitat, climate and soil conditions of these locations.
Tiger beetles are known to
prefer riverine habitats because of close proximity to food and water
resources, safety from predators and reasonably low human disturbance (Bhargav
& Uniyal 2008). Shade of the
habitat is used as an oviposition cue for many species and clusters of larval
burrows were found near the bases of plants in Cicindela cursitans Le Conte, of Ohio
Valley, United States (Brust et al. 2005).
Further, their activity and
density are greatly influenced by daily temperature patterns (Schultz 1983),
and adult tiger beetles are known to maintain internal body temperatures that
are just below their lethal limits of 390C (Pearson et al. 2006). A ground temperature ranging from 32-330C is known to be suitable for the activity and viability of tiger
beetle populations, and a temperature of 34-35 0C determined the
greatest number of matings in Cicindela (Cephalota) circumdata leonschaeferi Cassola (Eusebi et
al. 1989).
Tiger beetles prefer sandy
soils with minimal vegetation, where periodic disturbance by wind and water
removes encroaching vegetation (Warren & Buttner 2008). Colour patterns of
adult tiger beetle species closely match the texture and hue of the soil substrate
on which the species occurs (Pearson & Vogler 2001), and matching the body
colour with that of the soil plays an important role in predator evasion by
reducing discovery (Morgan et al. 2000).
The habitats at Maha Oya
(Dehi Owita) and Handapangoda were both riparian with sparse vegetation that
consisted of shrubs. In
Handapangoda, ferns and bamboo were found on the stream banks providing shade
to the habitat while a similar habitat was also found at Maha Oya. Ground temperature were similar at both
sites and were at 320C, while the soil temperature was 280C
at Maha Oya, and 270C at Handapangoda. Both river banks consisted of soil with a sandy texture that
was yellowish-brown at Maha Oya and dark yellowish-brown at Handapangoda. The colour of soil at both sites
closely matched the colour of beetles, that were metallic brown dorsally with
yellow-white maculations.
As C. (I.) willeyi is an endemic species
with limited distribution and is consequently susceptible to local extinction,
it is important that these habitats are identified and protected. At present, 99.5% of its distributional
range has declined (Table 6). We
believe that the most important conservation priority of a country is the
protection of areas which house large numbers of endemic species and
communities found nowhere else in the world. This study presents first evidence and importance of Maha
Oya (Dehi Owita) and Handapangoda locations, as harbouring the only populations
of an endemic species of tiger beetles. In view of the current human use of all of these sites and habitats and
the development pressures exerted on the wet zone of the country, conservation
of these sites are essential for future survival of this species.
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