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Eight New Microsatellite Loci of the Western Palearctic Hyles Euphorbiae Complex (Lepidoptera, Sphingidae)

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Eight New Microsatellite Loci of the Western Palearctic Hyles Euphorbiae Complex (Lepidoptera, Sphingidae) Ann. Zool. Fennici 48: 142–146 ISSN 0003-455X (print), ISSN 1797-2450 (online) Helsinki 30 June 2011 © Finnish Zoological and Botanical Publishing Board 2011 Eight new microsatellite loci of the Western Palearctic Hyles euphorbiae complex (Lepidoptera, Sphingidae) Michael B. Mende1,2,*, Heiko Stuckas1 & Anna K. Hundsdoerfer1,2 1) Museum of Zoology (Museum für Tierkunde), Senckenberg Natural History Collections Dresden, Königsbrücker Landstr. 159, D-01109 Dresden, Germany (*corresponding author; e-mail: [email protected]) 2) Biodiversity and Climate Research Centre (BiK-F), Senckenberganlage 25, D-60325 Frankfurt am Main, Germany Received 1 Nov. 2010, revised version received 23 Feb. 2011, accepted 28 Feb. 2011 Mende, M. B., Stuckas, H. & Hundsdoerfer, A. K. 2011: Eight new microsatellite loci of the Western Palearctic Hyles euphorbiae complex (Lepidoptera, Sphingidae). — Ann. Zool. Fennici 48: 142–146. We describe eight new microsatellite loci for hawkmoths of the Hyles euphorbiae complex. They are polymorphic (except for one locus in one population) with 2–18 alleles per locus, an expected heterozygosity between 0.14 and 0.94, and an observed heterozygosity between 0.10 and 0.75. As typical for Lepidoptera, the yield of new loci was low due to the presence of microsatellite gene families and variable flanking regions. These microsatellites provide informative results in population studies of the West Palearctic H. euphorbiae complex since cross amplification for H. euphorbiae and H. tithymali was successful for all but one locus. The taxa of the Hyles euphorbiae complex as nuclear as well as mitochondrial sequence mark- well as more distantly related species are known ers. This indicates young lineages in statu nas- to readily hybridize in nature (Harbich 1975, cendi and/or reintegration of incompletely sepa- 1976a, 1976b, see overview in Hundsdoerfer et rated lineages which makes the species com- al. 2005b). A natural hybrid population between plex especially interesting for population genetic the two main lineages, namely the North African studies. To reveal the patterns of natural hybridi- H. tithymali (Boisduval 1832) and the Euro- zation between the lineages a co-dominant geno- pean H. euphorbiae (Linnaeus 1758), is postu- type marker system is essential. lated for Malta based on morphological (Har- Microsatellite markers have been reported to bich 1989, 2009) and mitochondrial sequence be challenging in identification and application data (Hundsdoerfer et al. 2005a, 2005b, 2009b). for lepidopteran species (Meglécz et al. 2004). Overall, the reproductive plasticity is reflected They are rare within lepidopteran genomes and in fuzzy species delimitations caused by a high difficult to amplify (Meglécz & Solignac 1998). variability of the few applicable morphologi- Lepidopteran microsatellites are associated with cal characters, especially larval colour patterns flanking regions characterized by repetitive (Hundsdoerfer et al. 2011). Hundsdoerfer et al. sequences (Meglécz et al. 2004), a high inci- (2005a, 2005b, 2009b, and unpubl. data) and dence of single nucleotide polymorphisms and Hundsdoerfer and Wink (2006) found consider- indels (Hundsdoerfer et al. 2009a) and mobile able incongruences between morphology and elements leading to multiple copies of a locus ANN. ZooL. FENNIcI Vol. 48 • Eight new microsatellite loci of the Hyles euphorbiae complex 143 (microsatellite families, Zhang 2004, Hunds- at the 5´ end and approximately 10–30 ng DNA doerfer et al. 2009a). These properties can result template. The PCR comprises 38 cycles of dena- in a deficit of heterozygotes due to the presence turation at 94 °C for 30 s (5 min for the first of null alleles (Meglécz et al. 2004). Neverthe- cycle), primer annealing at 56 °C for the first 30 less, microsatellites can be used for lepidopter- cycles and 53 °C for the last 8 cycles for 45 s, ans with proper attention being paid to these and extension at 72 °C for 45 s (10 min for final issues (Hundsdoerfer et al. 2009a). elongation). Primers for loci Hti63 and Hti67 DNA was isolated from larvae of Hyles performed unsufficiently with dye-label ing after euphorbiae (Iberia) and Hyles tithymali (Canary the protocol of Schuelke (2000), but worked well Islands) using the innuPREP DNA Mini Kit as directly labeled forward primers (0.4 µM) (Analytik Jena, Germany) to create a DNA with a modified protocol (94 °C for 3 min, 39 library for each species (performed by GENter- cycles of 30 s at 94 °C, 30 s at 59 °C for Hti63 prise, Mainz, Germany): DNA was sheared by or 57 °C for Hti67 respectively and 1 min at nebulization. After electrophoretic fractionation 72 °C and final 20 min at 72 °C). Fragment fragments of two size ranges (1.5 to 2.5 kb and size was determined on an ABI3130 automatic > 2.5 kb) were excised and regained by electro- sequencer using an internal size standard (Liz elution, ligated into SmaI-cut, dephosphorylated 600, ABI) and the software GeneMapper (ver. 4, pUC19 vectors and transformed into Escherichia ABI). coli host cells (tribe DH10B). Approximately The eight loci were polymorphic in all four 16 000 colonies for H. tithymali and H. euphor- tested populations of both species (2–18 alleles, biae each were screened for the microsatellite Table 1), except for locus Heu76 which was not motives GA, CA, AAT, GGC, AAG and ATG by reliably PCR amplified in H. tithymali and locus hybridization of colony filters with correspond- Hti63 in which one allele got fixed in the Bul- ing oligo probes. garian population of H. euphorbiae (Table 1). One hundred and ninety two positive clones Although this respective allele is missing the (96 per species) were sequenced (GENterprise) microsatellite we argue that locus Hti63 still has of which 146 (85 for H. euphorbiae and 61 for some population genetical value because of a H. tithymali) contained microsatellite repeats. cline with higher frequencies of alleles contain- The flanking regions of 59 clones in total were ing a repeat motif in more southwestern popula- appropriate to design primer pairs (GENter- tions. Significant differences between expected prise). Only eight loci (14%) for H. euphor- heterozygosity (ranging from 0.14 and 0.94) and biae (and seven for H. tithymali respectively) observed heterozygosity (ranging from 0.10 and were reliably PCR amplified and showed to be 0.75) were found at five loci (Table 1, calculated polymorphic without indication of microsatel- and tested using Arlequin ver. 3.1.1, Excoffier lite families (multiple alleles per individual). et al. 2005). The heterozygote deficit at these These eight loci (Table 1) of which five were loci can be explained by the presence of null originally found in H. tithymali and three in H. alleles (Table 1; inferred by the software Micro- euphorbiae were used to genotype 20 non-sib- Checker ver. 2.2.3, van Oosterhout et al. 2004). ling individuals each from two populations of H. A significant heterozygote excess was found at euphorbiae (southeast Bulgaria, BUL; northeast one locus in one population (inferred by Gene- Spain, ESP) and two populations of H. tithymali pop on the web ver. 4.0: http://genepop.curtin. (Gran Canaria, GC; La Palma, LP). PCR was edu.au/, Raymond & Rousset 1995, Rousset performed in a volume of 20 µl following the 2008; Table 1). The definition of populations (i.e. protocol of Schuelke (2000) containing 1 U Taq pooling of specimens from different localities) polymerase (Bioron, Ludwigs hafen, Germany), might also be the reason for Hardy-Weinberg PCR reaction buffer with a final concentration disequilibrium in some cases. Furthermore, these of 2.5 mM MgCl2 (Bioron), 0.2 mM of each problems could result from the peculiar nature of dNTP (Fermentas, St. Leon-Rot, Germany), 0.4 H. euphorbiae for which the hybrid influx from µM FAM labeled M13-Primer, 0.4 µM reverse other species of the genus has been hypothesized Primer, 0.1 µM forward Primer with M13 tail (Harbich 1976a). Linkage disequilibrium was 144 Table 1. characteristics of eight microsatellite loci studied in Hyles euphorbiae from Europe (Bulgaria & Spain) and H. tithymali from the canary Islands (Gran canaria & La Palma). Abbreviations: Pop = population, BUL = population from Southeast Bulgaria, ESP = population from Northeast Spain, Gc = population from Gran canaria, LP = population from La Palma, n number of specimens analyzed, HE expected heterozygosity, Ho observed heterozygosity, pHWE probability that population is in Hardy Weinberg equilibrium. b Primer name and sequence Repeat GenBank Species Pop n No. of Allele size HE Ho pHWE Null sequencea acc. no. alleles allele bias X Hti62F 5´-GAGAAcGTcGGcAAATAATcGT-3´ (Ac)10 JF412019 H. euphorbiae BUL 20 8 369–427 0.86 0.10 < 0.001 Yes Hti62R 5´-GTGcAGATGGcAGTcTTATcG-3´ H. euphorbiae X ESP 18 18 359–432 0.93 0.50 < 0.001 Yes H. tithymali Gc 20 5 359–378 0.58 0.40 0.224 No H. tithymali LP 18 4 363–378 0.64 0.72 0.035 No X Hti63F 5´-TATATAGAATGGAGGGTGAcATTG-3´ (Tc)5 JF412020 H. euphorbiae BUL 20 1 269 – – – – Hti63R 5´-AAGcAAccTAATATTcAAcGccA-3´ H. euphorbiae X ESP 20 3 268–283 0.14 0.10 0.071 No H. tithymali Gc 20 2 269–283 0.47 0.70* 0.041 No H. tithymali LP 19 3 269–283 0.53 0.42 0.349 No X Hti65F 5´-cTGGcTAAAGTGTTcATGAGcT-3´ (Ac)5(cA)2 JF412021 H. euphorbiae BUL 20 5 313–322 0.64 0.75 0.214 No Hti65R 5´-cTcAATATcTGTTcGTAcTcGAG-3´ H. euphorbiae X ESP 20 8 282–324 0.63 0.45 0.136 No H. tithymali Gc 20 4 239–322 0.64 0.60 0.055 No H. tithymali LP 20 3 239–322 0.66 0.65 0.378 No X Hti66F 5´-AAGATcTTGcccTGGTTGAAc-3´ (cA)8 JF412022 H. euphorbiae BUL 20 2 250–260 0.45 0.15 0.005 Yes Hti66R 5´-AccTcTAcAATcGTGGAGcc-3´ H.
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