J. Limnol., 2016; 75(3): 571-580 ORIGINAL ARTICLE DOI: 10.4081/jlimnol.2016.1476 This work is licensed under a Creative Commons Attribution-NonCommercial 4.0 International License (CC BY-NC 4.0). Biogeography of the water flea Daphnia O. F. Müller (Crustacea: Branchiopoda: Anomopoda) on the Indian subcontinent Sameer M. PADHYE,1* Alexey A. KOTOV,2,3 Neelesh DAHANUKAR,4 Henri J. DUMONT5,6 1Wildlife Information Liaison Development Society, Coimbatore 641035, India; 2A. N. Severtsov Institute of Ecology and Evolution, Leninsky Prospect 33, Moscow 119071, Russia; 3Kazan Federal University, Kremlevskaya 18, Kazan 420000, Russia; 4Indian Institute of Science Education and Research, G1 Block, Dr. Homi Bhabha Road, Pashan, Pune 411008, India; 5Biology Department, Gent University, Ghent, Belgium; 6Department of Ecology and Hydrobiology, Jinan University, Guangzhou, China *Corresponding author: [email protected] ABSTRACT Studies on Daphnia distribution in Indian subcontinent have been few and regionally restricted despite Daphnia being by far the most studied cladoceran. We here present a first biogeographical assessment of the genus on the Indian subcontinent (Afghanistan, Pakistan, India, Nepal, Bhutan, Bangladesh and Sri Lanka). We collected all pertinent literature and considered nineteen bioclimatic variables along with latitude, longitude, and altitude for statistical analysis of factors governing distribution in space. Significant variables (determined by Kruskal Wallis test) were tested by nonparametric multivariate analysis of variance (PERMANOVA) to clarify whether Daphnia species had specific environmental requirements. Canonical correspondence analysis was used to understandonly how environmental variables affected distribution. Eight Daphnia (Ctenodaphnia) and 4 Daphnia s.str. occurred at 100 different localities. The variables temperature, altitude and latitude differed among species and so did their bio-climatic requirements. Daphnia distribution responded positively to altitude and negatively to a decrease in latitude and temperature. We confirm the existence of three complexes of Daphnia in the Indian subcontinent: i) widely distributed species and species complexes; ii) high altitude endemics; anduse iii) low latitude D. (Ctenodaphnia) species. Key words: Ctenodaphnia; Daphnia magna; Daphnia pulex; Himalayas; oriental zone; Western Ghats. Received: March 2016. Accepted: June 2016. INTRODUCTION (Ishida and Taylor, 2007; Mergeay et al., 2008; Adamow- icz et al., 2009) but little information is available on the The Cladocera (Crustacea: Branchiopoda) constitute Indian subcontinent. Yet, this region is interesting for bio- an ancient group of primarily freshwater crustaceans geography because of i) a complex geological history; ii) whose major differentiation may have occurred before the a wide range of altitudes; and iii) a strong seasonality with Permian (Kotov and Taylor, 2011). Until the Late Ceno- a distinct rainfall season (Monsoon) (Mani, 1974; Briggs, zoic, the Cladocera (and other Branchiopoda) are believed 2003). This has allowed the formation of various biomes, to have been much species-richer than today (Kerfoot and ranging from tropical to Alpine forests and deserts (Mani, Lynch, 1987; Korovchinsky, 2006). As a result of climatic 1974) resulting in a combination of endemic, Oriental, changes in the Tertiary, maximum cladoceran species Palaearctic as well as Gondwanan biota (Gower et al., richness is now concentratedNon-commercial in the subtropical zone of 2002; Whatley and Bajpai, 2006; Koehler and Glaubrecht, both hemispheres and at higher elevations within the true 2007; Kulkarni and Pai, 2016). Their presence has been tropics (Korovchinsky, 2006). explained by hypotheses like the ‘Indian raft’ and ‘Out of Daphnia (Anomopoda: Daphniidae) is among the Asia into India’ (Chatterjee and Scotese, 1999; Karanth, best-studied freshwater organisms (Lampert, 2011). It has 2003), but relevance of such scenarios for passively dis- an origin estimated at more than 200 Myr (Taylor et al., persed zooplankton is uncertain. 1996; Kotov and Taylor 2010, 2011). Currently the genus Here, our aim was to analyze Daphnia distribution on is classified into three sub-genera (Adamowicz et al., the Indian sub-continent, discuss associations between 2009) with 361 formally described species in toto of species and environmental variables, and evaluate the rel- which c. 24% are considered valid (Kotov, 2015). Among ative importance of selected climatic variables in eluci- them, only few inhabit the lowland equatorial zone (Fer- dating the distributional patterns observed. nando, 1980; Fernando and Kanduru,1984; Benzie. 2005), with species occurring only above particular altitudes METHODS (Dumont, 1980; Green, 1995). Study area The biogeographical patterns observed in many Daph- nia species in the Neotropics, Neartic, Paleartic and Afro- Our study area (Fig. 1) extends from Afghanistan and tropical regions have been assessed relatively adequately Pakistan to the west, India in the centre, Nepal and Bhutan 572 S.M. Padhye et al. to the east and Sri Lanka to the south (between 38°N to in peninsular region of the subcontinent reaches nearly 4°N latitudes and 60°E to 98° E longitudes). Geographi- 2600 m asl. The southernmost region of the subcontinent, cally, this region includes the Hindukush in the northwest, viz. Sri Lanka, has an altitudinal range of 0-2500 m. Himalayas from north till north-east; Indo-Gangetic plains Most of the Indian subcontinent has a monsoon cli- on the Himalayan slopes, a central plateau region (like mate, with rains for 3-4 months, usually from June till Deccan plateau) flanked by the more continuous Western September/October. Part of the southern region and the and more fragmented Eastern Ghats, and a peninsular re- north-eastern zone also receive rains from the retreating gion bordered by a narrow coastline (Mani, 1974). monsoon in November and December. An altitudinal gradient is seen as we move down along the latitude. The altitude of the Himalayas ranges from Data collection and GIS analysis 6600 m asl in northern and northwestern part (Tibetan Hi- malayas) to 900-1200 m asl in the more southern Sivalik Data were gathered through mining of the literature ranges. The Central Indian subcontinent (plateau region) published on Daphnia faunistics, taxonomy and biology. has an altitudinal range of 350-900 m asl, the highest peak We used a conservative approach in selecting literature only use Non-commercial Fig. 1. Indian subcontinent considered for the study along with distribution of study sites (circles represent each locality). Biogeography of Daphnia in the Indian subcontinent 573 on Daphnia occurrence because a large number of Daph- Diversity and taxonomy nia records from the sub-continent are dubious (Chatterjee Twelve species of Daphnia have been reported from the et al., 2013). We only accepted records having at least region (few comprising of presumable species groups) some brief description of the animals with illustrations (Tab. 3), of which eight belong to the subgenus D. (Cten- making possible a check of the identification of the cited species (for all records, see Supplementary Tab. 1). No reliable records of Daphnia could be obtained from Bangladesh and only a single report of D. tibetana was Tab. 1. Names of the BIOCLIM variables considered for the study. available from Bhutan (H. Dumont, pers. obs.). Latitude and longitude data on each locality were ob- BIOCLIM Description tained from Google Earth portal. A set of 19 bioclimatic Bio1 Annual mean temperature variables and altitudes were extracted from the BIOCLIM Bio2 Mean diurnal range (mean of monthly) (max temp - min temp) dataset (http://www.worldclim.org; Hijmans et al., 2005) Bio3 Isothermality - (Bio2/Bio7) (*100) at ten-minute spatial resolution (for names see Tab. 1). Bio4 Temperature seasonality (standard deviation *100) Broad scale resolution was used as many of the localities Bio5 Max temperature of warmest month Bio6 Min temperature of coldest month given were quite nonspecific (Ex. Kolkata as a locality). Bio7 Temperature annual range (Bio5-Bio6) Bio8 Mean temperature of wettest quarter Data analysis Bio9 Mean temperature of driest quarter Environmental variables including latitude, longitude, Bio10 Mean temperatureonly of warmest quarter altitude and the 19 bioclimatic variables were considered Bio11 Mean temperature of coldest quarter Bio12 Annual precipitation for statistical analysis. Univariate normality of each vari- Bio13 Precipitation of wettest month able was checked using Shapiro-Wilk test. Since all vari- Bio14 use Precipitation of driest month ables proved to be non-normal, variables that were Bio15 Precipitation seasonality (coefficient of variation) significantly different among Daphnia species were iden- Bio16 Precipitation of wettest quarter tified using non-parametric Kruskal-Wallis one-way Bio17 Precipitation of driest quarter analysis of variance. As multiple tests were performed, Bio18 Precipitation of warmest quarter the family-wise error rate was corrected using Bonferroni Bio19 Precipitation of coldest quarter correction. Significant variables were then used in analyzing Daphnia distribution. The null hypothesis that the data is Tab. 2. Kruskal-Wallis one-way analysis of variance test for the multivariate normal was checked using an omnibus test environmental variables across different species. Underlined P (Doornik and Hansen, 2008). Since the data was not mul- values are significant after Bonferroni