MAJOR ARTICLE

TAPROBANICA, ISSN 1800–427X. August, 2015. Vol. 07, No. 04: pp. 224–234. © Research Center for Climate Change, University of Indonesia, Depok, Indonesia www.taprobanica.org

AN INVERTEBRATE PERSPECTIVE TO HUTCHINSON’S RATIO USING CO-OCCURRING TIGER (COLEOPTERA: CICINDELIDAE) ASSEMBLAGES

Section Editor: Jörg Gebert Submitted: 31 December 2014, Accepted: 27 July 2015

Chandima Deepani Dangalle1, 2 and Nirmalie Pallewatta1

1 Department of Zoology, Faculty of Science, University of Colombo, Cumaratunga Munidasa Mawatha, Colombo 03, P.O. Box 00300, Sri Lanka; 2 E-mail: [email protected]

Abstract Tiger co-occur for cryptic advantage, interspecific hybridization, reproductive synchrony and thermoregulation. Successful co-occurrence relies on the absence of competition and Hutchinson (1959) has proposed body size ratios for co-occurrence of bird and mammal taxa. The present study calculates body size ratios for co-occurring assemblages and investigates similarities with ratios for co-occurring birds and mammals. Ratios obtained using measurements of different morphological characters are considered and compared. Association of body size ratios of co- occurring tiger beetles with habitat type and number of species in co-occurring tiger beetle assemblage are determined. The study revealed a body weight ratio and body length ratio for co- occurring tiger beetles that was similar to Hutchinson’s ratios for birds and mammals. Different ratios were obtained when using different morphological parameters. Body size ratios inferred using mandible lengths differed significantly according to habitat type. Further, relatively higher values were observed for assemblages occupying terrestrial habitats while lower values were found for assemblages of aquatic habitats.

Keywords: body size, competition, morphology, habitat type, Sri Lanka

Introduction Many “co-occur” in multispecies in communities invaded by other species assemblages in various ecosystems. Biotic (Sanders et al., 2003) and the rich diversity of interactions such as beneficial hybridization foods available in an ecosystem (Thorington & (Chunco et al., 2012), heterospecific attraction Ferrell, 2006) are known to foster co-occurrence where heterospecifics indicate the suitability of a amongst many taxa. However, co-occurrence habitat (Sebastian-Gonzalez et al., 2010), leads to competition that results in loss of aggregation effect of native species occurring species (Westman & Savolainen, 2001), juvenile

224 TAPROBANICA VOL. 07: NO. 04 AN INVERTEBRATE PERSPECTIVE TO HUTCHINSON’S RATIO bottlenecks (Guruge & Amarasinghe, 2008), temperature, soil characteristics and oviposition non-detection of certain species that leads to choice (Ganeshaiah & Belavadi, 1986; Hoback conservation issues (Bailey et al., 2009) and et al., 2000; Woodcock et al., 2010). Further, predation (Taggart et al., 2005). Therefore, co-existence as a communal roost has been competition and other unfavourable conditions related with cryptic advantage, thermoregulation should be absent for successful co-occurrence to and synchrony of reproduction and dispersal prevail. In 1959, Hutchinson suggested that for (Bhargav & Uniyal, 2008). Interspecific ecologically similar species to co-occur a hybridization in co-occurring tiger beetle species constant ratio between body sizes must persist (Brust et al., 2012), as well as reproductive (Hutchinson, 1959; Simberloff & Boecklen, isolation in some cases of sympatric species 1981; Weins, 1982; Eadie et al., 1987). By the (Tigreros & Kattan, 2008) have also been examination of birds and mammals he suggested addressed. Satoh et al. (2003) investigated body an average ratio (larger/smaller) of 1.3 for body size differences in co-occurring coastal tiger length and 2.0 for body weight (Weins, 1982; beetle species assemblages in Japan. The study Eadie et al., 1987). Many studies further considered the head width and mandible length demonstrated the use of other morphological of species and relative overlaps between species measurements – skull length, head width, bill were calculated. However, the ratios proposed length, jaw length, carapace width, ovipositor by Hutchinson (1959) have not been calculated length, proboscis length, wing length – for for co-occurring tiger beetle species and whether calculation of this average ratio (Weins, 1982). the patterns adhere to the values envisioned by However, as the measurements of different Hutchinson has not been considered. If ratios morphological characters produced different adhere to the proposed ratios they can be used to ratios, and in some cases different size-sequence support Hutchinson’s ratio from an invertebrate structuring for similar morphological characters perspective and further, as evidence for absence in different species, the ecological relevance of of competition. the characters used need to be considered (Weins, 1982). Therefore, in the present study we calculated body size ratios for co-occurring tiger beetle Tiger beetles (Coleoptera, Cicindelidae) are assemblages using morphometric data that have omnivorous predators that occur in temperate been recorded in previous studies and current and tropical environments. Many adult tiger study conducted in Sri Lanka. Ratios obtained beetles co-occur and are found as sympatric using measurements of different morphological assemblages of two to fifteen species in a range characters were considered and compared. of habitats (Cornelisse & Hafernik, 2009). Ten Association of body size ratios of co-occurring species of tiger beetles co-occur on flats and tiger beetles with habitat type and association of stream banks of the Eastern Nebraska salt marsh body size ratios with number of species in co- of which one species is a threatened in the occurring tiger beetle assemblage were United States (Hoback et al., 2001). Eight determined. Further, co-occurring tiger beetle species co-exist on the floodplains of the Tedori populations of Sri Lanka were investigated and River System of Japan (Satoh et al., 2006). species assemblages, locations and habitat types Three species co-exist as a communal roost on recorded. The study aimed to detect a pattern in plant species of Simbalbara Wildlife Sanctuary, size relationships of co-occurring tiger beetle Shivalik Hills, India (Bhargav & Uniyal, 2008). species and whether body size ratios can be used Four species of tiger beetles are found on the as an index to suggest co-existence of species. river beds of Arkavathy and Cauvery river, Karnataka, India (Ganeshaiah & Belavadi, Materials and methods 1986), and two species occur sympatrically in Compilation of data from previous studies: the mountain habitats of Colombia (Tigreros & Data on co-occurring tiger beetle species was Kattan, 2008). In Japan, nine species of tiger obtained from nine studies conducted from 1979 beetle co-occur in estuarine seashore and sand to 2010. Co-occurring species, locations, habitat dunes along the coast (Satoh et al., 2003). types and body size ratios obtained from morphometric measurements were recorded for Studies of co-occurring tiger beetles have tiger beetle assemblages reported in the studies concentrated on niche partitioning of species (Appendix 1). Body size ratios (larger species according to preferences for prey, vegetation, over the smaller one) for co-occurring species

225 TAPROBANICA VOL. 07: NO. 04 DANGALLE & PALLEWATTA, 2015 were calculated using morphometric data: body Mandible length was estimated by measuring the weight; body length, from the frons of head to distance from the articulation point to the tip of the elytral apex; head width, maximum width the left mandible. Broken and worn out including eyes; mandible gape, the space mandibles were disregarded (Pearson & Juliano, between open mandibles; mandible length, 1991). Measurements of both body length and distance between articulation point and the tip of mandible length were taken using a dissecting the mandible; and elytral length, distance microscope (Nikon Corporation SE, Japan) with between base of scutellum and apex of elytra. an eyepiece graticule (Nikon, Tokyo, Japan) that When an assemblage consisted of more than two was calibrated by an objective micrometer species, ratios were calculated by considering (Olympus, Japan). the association of each species with every other species. Data analysis: Body size ratios obtained from

different morphological characters were Investigation and collection of tiger beetles compared using One-Way Analysis of Variance from Sri Lanka: Investigations for tiger beetles and Tukey’s Multiple Comparison method of the were conducted in ninety-four locations of Sri Minitab 16.0 statistical software package. Lanka from May 2002 to December 2006. Association between habitat type and mandible Coastal areas, river banks, reservoir banks, length ratios of co-occurring tiger beetle agricultural lands, marsh lands and urban areas assemblages, and association between number of were investigated in the wet, intermediate and co-occurring species in an assemblage and dry zones of the country. When beetles were mandible length ratios were analyzed using the encountered a sample of three to five beetles of same test and software. The sample size for each species was collected using a standard mandible length ratios was large when insect net and preserved in 70% alcohol for comparing with ratios obtained for other identification. Permission to make collections of morphological data. Therefore, they were tiger beetles was obtained from the Department utilised in the analysis for demonstrating the of Wildlife Conservation, Sri Lanka. association between body size ratios and habitat Identification of tiger beetles: Tiger beetles type and association between body size ratios were identified using keys for Cicindela (sensu and number of co-occurring species in an lato) of the Indian subcontinent (Acciavatti & assemblage. Pearson, 1989) and descriptions of Horn (1904), Fowler (1912) and Naviaux (1984). Results Identifications were confirmed by comparing the Information obtained from previous studies specimens with type specimens available in the and current study: Data consisted of a total of National Museum of Colombo, Sri Lanka and sixty-four tiger beetle assemblages of which British Natural History Museum of London, fifty-eight assemblages were from previous United Kingdom. Nomenclature is based upon studies and six assemblages were from Sri Wiesner (1992) except for the use of Calomera Lanka. Each assemblage consisted of two to Motschulsky, 1862 instead of Lophyridia, based twelve tiger beetle species. Co-occurring upon Lorenz (2005). assemblages were recorded from five distinct

habitat types – Coastal ( sand dunes, salt flats, Measurement of morphological parameters of sea shores, beaches and bays), Reservoir (Pond tiger beetles: Co-occurring tiger beetle edges), Grassland, Forest (open forests, rain assemblages were identified and the body forests) and River (creeks, water canals, weight, body length and mandible length were floodplains). Tiger beetle assemblages of Sri measured and recorded for each specimen. Body Lanka were recorded from reservoir and river weight was estimated by weighing each beetle to bank habitats, and consisted of two to three the nearest mg on an analytical balance (Chyo species accounting for a total of seven species JL180, Chyo Balance Corp., Japan). Body (Appendix 1). length was estimated by measuring the distance from the frons of the head to the elytral apex when the head was in the normal feeding Body size ratios of co-occurring tiger beetle position. Caudal spines on the elytral apex were assemblages: Three hundred and fifty-eight disregarded (Acciavatti & Pearson, 1989). (358) body size ratios were calculated for the

226 TAPROBANICA VOL. 07: NO. 04 AN INVERTEBRATE PERSPECTIVE TO HUTCHINSON’S RATIO sixty-four assemblages of co-occurring tiger largest ratios were seen in assemblages beetles (Appendix 1). Body size ratios differed consisting of 6 species (p <0.01, F = 8.43) according to the morphometric parameter (Table 3). considered. Ratios obtained using body length, head width and mandible lengths were not statistically different, while body weight ratio was not statistically different to mandible gape ratio. A significant statistical difference was evident between body length, head width, mandible length ratios and body weight, mandible gape ratios (p <0.01, F = 10.32) (Table 1). Figure 1: Mandible length ratios of co-occurring Table 1: Body size ratios of co-occurring tiger beetle tiger beetle assemblages in different habitat types assemblages; means sharing a common letter(s) are not significantly different according to Tukey’s Table 3: Relationship between mandible length Multiple Comparison test. ratios and number of co-occurring tiger beetle Morphometric character Body size ratio species; means sharing a common letter(s) are not Body weight (n=10) 1.83a ±1.1 significantly different according to Tukey’s Multiple Mandible gape (n=21) 1.85a ±0.7 Comparison test. b Number of species in Body length (n=15) 1.26 ±0.2 Mandible length ratio b an assemblage Head width (n=51) 1.26 ±0.2 a b Two (n=24) 1.38 ±0.05 Mandible length (n=260) 1.37 ±0.3 b c Three (n=51) 1.52 ±0.04 Elytral length (n=1) 1.04 Four (n=66) 1.38a ±0.04 Five (n=40) 1.30c ±0.05 Mandible length ratios and habitat type: Six (n=15) 1.67d ±0.13 Significant differences were evident between Twelve (n=66) 1.22c ±0.03 mandible length ratios of co-occurring tiger beetles in different habitat types. Ratios for Discussion terrestrial habitat types (grasslands, forests) were The analysis of body size ratios of co-occurring significantly larger than the ratios for aquatic species has been a major focus in evolutionary habitat types (reservoir, riverine, coastal). The ecology. Hutchinson concluded that this value largest mandible length ratios were obtained for could be used as an indication of the kind of tiger beetle assemblages of grassland habitats, difference necessary to permit two species to co- while the smallest ratios occurred in species of occur in different niches but at the same level of reservoir habitats (p <0.01, F=11.00) (Fig. 1; a food-web. A body size ratio of 1.3 for body Table 2). length and 2.0 for body weight was interpreted as the amount of separation necessary to permit Table 2: Mandible length ratios of co-occurring tiger beetle assemblages in different habitat types; means co-occurrence of species at the same tropic sharing a common letter(s) are not significantly level. Larger organisms preferring larger prey different according to Tukey’s Multiple Comparison relative to smaller prey makes it necessary that test. similar size species must be more widely Habitat type Mandible length ratio separated avoiding minimum size ratios, while Coastal (n=78) 1.39a ±0.03 large ratios in assemblages result in more severe Riverine (n=30) 1.40a ±0.05 competition amongst species (Weins, 1982). The Reservoir (n=76) 1.22b ±0.03 present study revealed an average body weight Grassland (n=26) 1.67c ±0.09 ratio of 1.83 and an average body length ratio of Forest (n=51) 1.42d ±0.05 1.26 for co-occurring tiger beetle assemblages. These ratios were closely similar to the ratios Mandible length ratios and number of co- suggested by Hutchinson (body weight = 2.0, occurring species: A clear relationship was not body length = 1.3) for co-occurring birds and evident between mandible length ratio and mammals. Therefore, it is possible that the number of tiger beetle species in an assemblage. values proposed by Hutchinson in 1959 for The smallest ratios were observed in vertebrate taxa may be applicable for assemblages consisting of 12 species, while the invertebrates as well. However, consideration of

227225 TAPROBANICA VOL. 07: NO. 04 & DANGALLE & PALLEWATTA, 2015 other co-occurring invertebrate taxa in addition occurring tiger beetle species was not evident as to tiger beetles is required to confirm this larger ratios occurred in assemblages of four and finding. Body size ratios of tiger beetles differed six species, and ratios less than 1.3 occurred in according to the morphological measurement assemblages of five and twelve species. under consideration and mandible gape ratios were statistically similar to body weight ratios When considering the co-occurring tiger beetles while head width and mandible length ratios of Sri Lanka species were found on large sandy were statistically similar to body length ratios. banks of river and reservoir habitat types. Thus, the present investigation infers that co- Calomera angulata (Fabricius, 1798) was the occurring tiger beetles have body size ratios of main species in sympatric assemblages and was 1.26 – 1.37 when using body length, head width found in five of the six associations. Further, and mandible length, while ratios of 1.83 – 1.85 recent investigations in May 2014 revealed C. are obtained when using body weight and angulata co-occurring with another tiger beetle mandible gape. species in a riverine habitat at Deduru Oya, Puttalam District. C. angulata has also been Co-occurring tiger beetle assemblages of recorded in co-occurring tiger beetle different habitat types demonstrated different assemblages of riverine habitats of India and mandible length ratios which were larger for coastal habitats of Japan. In Sri Lanka, it is the terrestrial habitat types (grasslands, forests) and main species of reservoir habitats (Dangalle et smaller for aquatic habitat types (rivers, coastal al., 2012) and according to the present areas, reservoirs). This illustrated that species investigation the main species of sympatric tiger with a wide range of sizes can be expected in beetle assemblages. terrestrial assemblages while species with a narrower range of sizes can be expected in Our study reveals that co-occurring tiger beetle aquatic assemblages. Body size of tiger beetles species display body weight and body length is specific to habitat type (Dangalle et al., 2013). ratios similar to the ratios proposed by Large species are specific to reservoir and Hutchinson (1959) for birds and mammals. coastal habitats, while small species occupy Ratios can be obtained using different riverine habitat types (Dangalle et al. 2013). morphological characters, and mandible gape Such associations can narrow the body size ratios of tiger beetles are similar to body weight ranges of taxa occupying a habitat and result in ratios, while mandible length and head width smaller body size ratios. The present study ratios are similar to body length ratios. Mandible revealed body size ratios for tiger beetle length ratios of co-occurring tiger beetle assemblages of reservoir habitats to be assemblages in terrestrial habitats are larger than significantly small. This is strongly possible due for tiger beetles of aquatic habitats, which may to the high significance of body size – habitat be due to the close association of body size of type relationship of tiger beetles recorded from tiger beetles with aquatic habitat types. Further reservoir habitats (Edirisinghe et al., 2014). investigations in terrestrial habitats are required However, the relationships of tiger beetle body for more accurate inferences. size with terrestrial habitats such as forests and grasslands have not been investigated. The large Acknowledgements body size ratios revealed in the present study We thank the Department of Wildlife indicates the possibility of species assemblages Conservation, Sri Lanka for providing permits with a wide range of body sizes for these habitat necessary for field work and sample collection; types. S. M. Henkanaththegedara (Longwood University, USA) for valuable comments. The According to Pearson and Vogler (2001), if study was funded by the National Science three or fewer species of tiger beetles co- Foundation of Sri Lanka, grant numbers occurred, the ratio of their mandible lengths is RG/2003/ZOO/01 and RG/2012/NRB/02. greater than 1.3. Similarly, in the present investigation a mandible length ratio greater Literature Cited than 1.3 occurred in tiger beetle assemblages Acciavatti, R. E. and D. L. Pearson, 1989. The consisting of two and three tiger beetle species. tiger beetle genus Cicindela (Coleoptera, Insecta) However, a consistent association between from the Indian subcontinent. Annals of the mandible length ratio and number of co- Carnegie Museum, 58: 77–355.

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Appendix I: Data for co-occurring tiger beetles

Body Body Mandible Head Elytral Mandible length Location Co-occurring species weight length gape width length Source ratio ratio ratio ratio ratio ratio

1.02, 1.16, 1.02, 1.06, 1.22, 1.04, 1.06, 1.88, 1.06, 1.14, 1.07, 1.19, Cicindela haemorrhagica 1.04, 1.04, 1.25, 1.06, Cicindela marutha 1.09, 1.92, 1.08, 1.17, Cicindela ocellata 1.04, 1.14, 1.23, 1.05, Cicindela pimeriana 1.12, 1.10, 1.62, 1.10, Pond edge, Cicindela praetextata 1.02, 1.24, 1.08, 1.22, Pearson & Sulphur Springs Cicindela punctulata 1.02, 1.04, 1.85, 1.04, Mury Valley, Cicindela sedecimpunctata 1.12, 1.08, 1.10, 1.30, (1979) Arizona, USA Cicindela tenuisignata 1.13, 1.99, 1.12, 1.21, Cicindela viridisticta 1.00, 1.18, 1.15, 1.54, Cicindela nevadica 1.16, 1.07, 1.30, 1.02, Cicindela nigrocoerulea 1.81, 1.02, 1.10, 1.11, Cicindela willistoni 1.78, 1.00, 1.08, 1.13, 1.78, 1.65, 2.00, 1.08, 1.13, 1.21

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Body Body Mandible Head Elytral Mandible length Location Co-occurring species weight length gape width length Source ratio ratio ratio ratio ratio ratio

Cicindela obsolete Grassland, Cicindela pulchra 1.02, 1.50, 1.36, 2.46, Pearson & Sulphur Springs Cicindela nigrocoerulea 2.25, 1.52, 1.38, 2.50, Mury Valley, Cicindela horni 2.29, 1.10, 1.65, 1.51, (1979) Arizona, USA Cicindela lemniscata 1.81, 1.66, 1.09 Cicindela debilis 1.17, Arkavathy, Cicindela cardoni 1.16, Ganeshaiah Cauvery River Cicindela cancellata 1.80, 1.25, 1.09, 1.63, 1.36, & Belavadi Junction, Cicindela sumatrensis 1.00, 1.77, 1.30 (1986) Karnataka, India Cicindela minuta 1.55, 1.54 Schultz & Carizzo Creek, Cicindela tranquebarica, 1.21 Hadley Arizona, USA Cicindela oregona (1987) Cicindela limbata Sand dunes, Fox Cicindela scutellaris 1.15, 1.29, 1.47, 1.12,

Valley, Canada Cicindela lengi 1.27, 1.13 Cicindela formosa Cicindela purpurea 1.01 Cicindela tranquebarica Open forest, Pennsylvania, Cicindela scutellaris Cicindela purpurea 1.01, 1.03, 1.05, 1.02, USA Cicindela patruela 1.04, 1.02 Cicindela tranquebarica Cicindela fulgida Cicindela togata 1.20, 1.24, 1.30, 1.30, Salt flat, New Cicindela circumpicta 1.03, 1.08, 1.08, 1.04, Mexico, USA Cicindela nevadica 1.05, 1.00 Cicindela willistoni Cicindela togata Cicindela circumpicta 1.15, 1.22, 1.25, 1.29, Salt flat, Kansas, Cicindela nevadica 1.06, 1.08, 1.11, 1.02, USA Cicindela fulgida 1.06, 1.03 Cicindela willistoni Cicindela lemniscata Pearson & Cicindela nigrocoerulea 1.63, 2.22, 1.36 Juliano Cicindela obsoleta (1991) Cicindela obsolete 1.26 Grassland, Cicindela horni Arizona, USA Cicindela debilis Cicindela horni 1.73, 2.19, 1.26 Cicindela obsoleta Cicindela lemniscata Cicindela nigrocoerulea 1.63, 2.35, 1.44 Cicindela pulchra Cicindela punctulata Permanent pond Cicindela ocellata 1.03, 1.17, 1.31, 1.13, edge, Arizona, Cicindela sedecimpunctata 1.27, 1.12 USA Cicindela haemorrhagica Temporary pond Cicindela marutha edge, Arizona, 1.00 Cicindela fulgoris USA Grassland, Cicindela lefroyi 1.31 Chandigarh, India Cicindela striatifrons Permanent pond Cicindela melancholica edge, 1.46 Cicindela minuta Chandigarh, India

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Body Body Mandible Head Elytral Mandible length Location Co-occurring species weight length gape width length Source ratio ratio ratio ratio ratio ratio

Cicindela albopunctata Cicindela subtilesignata 1.39, 1.21, 1.68 Sandy river bank, Cicindela intermedia Pearson & Chandigarh, India Sandy river bank, Cicindela agnate Juliano Cicindela venosa 1.12, 1.43, 1.49, 1.82, Chandigarh, India (1991) Cicindela angulata 1.28, 1.32, 1.62, 1.04, Cicindela chloris 1.27, 1.22 Cicindela plumigera Lowland rain Cicindela striolata forest, Karnataka, Cicindela collicia 1.74, 2.33, 1.34 India Cicindela fabriciana Cicindela dasiodes Cicindela fastidiosa 1.18, 1.84, 1.88, 2.28, Open forest, Cicindela bicolor 1.57, 1.60, 1.94, 1.02, Karnataka, India Cicindela calligaramma 1.24, 1.21 Cicindela aurofasciata Cicindela undulate Sandy river edge, Cicindela minuta 1.23, 1.35, 1.86, 1.10,

Karnataka, India Cicindela cancellata 1.51, 1.38 Cicindela angulata Cicindela belli Highland rain Cicindela striolata 1.71, 1.94, 2.14, 1.13, forest, Kerala, Cicindela hamiltoniana 1.25, 1.11 India Cicindela andrewesi Lowland rain Odontocheila iodopleura forest, Heredia, 1.56 Odontocheila nicaraguensis Costa Rica Lowland rain forest, Odontocheila iodopleura 1.36 Puntarenas, Costa Odontocheila chiriquina Rica Lowland rain Odontocheila salvini forest, B.C. 1.37 Pearson & Odontocheila chiriquina Island, Panama Juliano Lowland rain Odontocheila trilbyana (1991) forest, Napo, Odontocheila consobrina 1.39, 1.62, 1.17 Ecuador Odontocheila cayennensis Lowland rain Odontocheila iodopleura forest, Pichincha, 1.35 Odontocheila chiriquina Ecuador Lowland rain Odontocheila rufiscapsis forest, Ucayali, 1.33 Odontocheila cayennensis Peru Lowland rain Odontocheila egregia forest, Beni, Odontocheila spinipennis 1.08, 1.51, 1.40 Bolivia Odontocheila confusa Lowland-terra Odontocheila luridipes rain forest, Odontocheila annulicornis 1.21, 1.37, 1.14 Tambopata, Peru Odontocheila cayennensis Lowland- Odontocheila lacordairei floodplain rain Odontocheila cyanella 1.09, 1.34, 1.73, 1.23, forest, Odontocheila annulicornis 1.59, 1.29 Tambopata, Peru Odontocheila confusa Open forest Odontocheila annulicornis bamboo, 1.07 Odontocheila chrysis Tambopata, Peru Lowland rain Therates batesi forest, Borneo, 1.63 Therates spinipennis Indonesia

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Body Body Mandible Head Elytral Mandible length Location Co-occurring species weight length gape width length Source ratio ratio ratio ratio ratio ratio

Lowland rain Therates rathschildi forest, Sepik, Therates festivus 1.30, 2.05, 1.57 Pearson & Papua New Therates basalis Juliano Guinea Cicindela nilotica (1991) Sandy river bank, Cicindela regalis 1.54, 2.09, 1.35 Kenya Cicindela dongalensis Salt Marsh, Cicindela circumpicta Hoback et Eastern Nebraska, 2.02 Cicindela togata al. (2001) USA Phaeoxantha lindemannae Black Water 1.33, 1.48, Phaeoxantha bifasciata Flood Plains, 2.33, 1.11, Tetracha punctata Manaus, Brazil 1.75, 1.58 Phaeoxantha limata 1.44, 1.64, Zerm & Phaeoxantha klugii 1.94, 3.59, Adis Phaeoxantha aequinoctialis 3.28, 1.14, White Water (2001) Tetracha punctata 1.35, 2.50, Flood Plains, Tetracha spinosa 2.29, 1.18, Manaus, Brazil Cylindera suturalis 2.19, 2.00, Pentacomia cribata 1.85, 1.69, 1.09 1.01, Sea Shore, Lophyridia angulata 1.28, Tanega-shima Abroscelis anchoralis 1.32, 1.75, 1.85, 1.33, 1.28, Island, Cylindera elisae 1.41, 1.06 1.26, Kagoshima, Callytron yuasai 1.26, Japan 1.00 1.06, Sea Shore, Lophyridia angulata 1.11, Shibushi Bay, Myriochile speculifera 1.60, 1.47, 1.95, 1.09, 1.39,

Kagoshima, Abroscelis anchoralis 1.32, 1.22 1.05, Japan Cylindera elisae 1.25, 1.31 Lophyridia angulata 1.11, Sea shore, Hyuga, Abroscelis anchoralis 1.42, 1.74, 1.22 1.31, Miyazaki, Japan Cylindera elisae 1.17 Sand dunes, Myriochile speculifera 1.02, Kaseda, Lophyridia angulata 1.43, 1.78, 1.24 1.27, Kagoshima, Cylindera elisae 1.30 Japan Satoh et al. (2003) 1.08, Cicindela lewisii 1.48, Fukiage beach, Lophyridia angulata 1.18, 1.74, 1.94, 2.04, 1.55, Kagoshima, Cylindera elisae 2.29, 1.12 1.37, Japan Callytron inspecularis 1.44, 1.05 Sea shore, Sendai, Cicindela lewisii 1.95 1.52 Kagoshima, Callytron inspecularis Japan Sea shore, Ohzai, Cicindela lewisii 1.66 1.42 Ohita, Japan Cylindera elisae 1.09, Cicindela lewisii 1.38, Sea shore, Myriochile speculifera 1.32, 1.64, 1.71, 1.24, 1.48,

Nagasaki, Japan Callytron yuasai 1.30, 1.04 1.27, Cylindera elisae 1.35, 1.07

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Body Body Mandible Head Elytral Mandible length Location Co-occurring species weight length gape width length Source ratio ratio ratio ratio ratio ratio

Sea shore, Cicindela lewisii 1.63 1.41 Hiroshima, Japan Cylindera elisae Sea shore, Cicindela lewisii 1.66 1.45 Tokoshima, Japan Cylindera elisae Sea shore, Chaetodera laetescripta 1.52 1.47 Shizuoka, Japan Cylindera elisae Sea shore, Chaetodera laetescripta 1.58 1.52 Ibaraki, Japan Cylindera elisae Sea shore, Lophyridia angulata 1.76 1.30 Shimane, Japan Cylindera elisae Lophyridia angulata 1.14, Sand dune, Cicindela tranbaicalica 1.31, 1.77, 1.35 1.32, Tanegaike, Japan Cylindera elisae 1.15 Chaetodera laetescripta 1.05, Tottori dune, Lophyridia angulata 1.24, 1.43, 1.74 1.36, Tottori, Japan Cylindera elisae 1.30 Satoh et al. Lophyridia angulata 1.07, Sea shore, (2003) Abroscelis anchoralis 1.27, 1.75, 1.38 1.34, Ishikawa, Japan Cylindera elisae 1.25 1.07, Chaetodera laetescripta 1.15, Sea shore, Lophyridia angulata 1.20, 1.00, 1.52, 1.21, 1.48,

Niigata, Japan Cicindela transbaicalica 1.83, 1.52 1.07, Cylindera elisae 1.38, 1.29 Tigreros & Western Andean Pseudoxycheila confuse 1.04 Kattan, Range, Colombia Pseudoxycheila chaudoiri (2008) Beach ridge, Cicindela l. longilabris Woodcock Wapusk National Cicindela limbata 1.25 et al. Park, Manitoba hyperborea (2010) Water canal, Cylindera (Ifasina) willeyi Handapangoda, Cylindera (Ifasina) 1.04 1.08 1.00 Kalutara district, waterhousei Sri Lanka Cylindera (Oligoma) Devahuwa wewa, 4.17, 1.41, lacunose Dambulla, Sri 3.56, 1.45, Lophyra (Lophyra) catena Lanka 1.17 1.03 Calomera angulata Kandalama Myriochila (Monelica) Dangalle et wewa, Dambulla, fastidiosa 1.14 1.05 1.17 al. Sri Lanka Calomera angulata (unpublishe Tabbowa wewa, Myriochila (Monelica) d data) Karuwalagaswew fastidiosa 1.03 1.01 1.01 a, Sri Lanka Calomera angulata Ma Oya, Calomera cardoni Alawwa, Sri 1.77 1.25 1.19 Calomera angulata Lanka Nachchaduwa Myriochila (Monelica) wewa, fastidiosa 1.21 1.12 Anuradhapura, Calomera angulata Sri Lanka

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