Geographic and Biological Variation in the Ibero-African Psilogaster
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Josef J. de Freina et al., Geographic and biological variation in Psilogaster Geographic and biological variation in the Ibero-African Psilogaster Reichenbach, 1817 and its taxonomical and biogeographical implications (Lepidoptera: Lasiocampidae, Pinarinae) ● Josef J. de Freina, Yeray Monasterio León, Joan Carles Hinojosa & Werner Bruer Abstract. Populations of Psilogaster Re- species are listed, and the different col- 1885), presente en el norte de África, se ichenbach, 1817 of the Iberian Penin- our patterns of their mature larvae as confirma como buena especie tanto por sula and North Africa are examined well as their habitats are described and una diferencia clara de un mínimo del based on molecular data from the mito- illustrated. 3.8 % en los datos genéticos moleculares, chondrial cytochrome oxidase subunit I como por las características de sus larvas. gene (COI) as well as on external and Resumen. Variación geográfica y bioló- Se revisa el estatus de Diploma simulatrix internal morphological structures. Their gica en el género íbero-africano de La- Chrétien, 1910 syn. rev. quedando como distribution and range of variability are siocámpidos Psilogaster Reichenbach, una forma de P. algeriensis. Se designa discussed. Psilogaster species are poly- 1817 y sus implicaciones biogeográficas un neotipo hembra para P. loti loti, que morphic in their colour and forewing y taxonómicas (Lepidoptera: Lasiocamp- queda depositado en el MWM (más tarde pattern. This has led to the description idae, Pinarinae). Se estudian las pobla- estará en la Colección Zoológica Estatal of several invalid species, as modifica- ciones de Psilogaster Reichenbach, 1817 de Múnic). Se enumeran las plantas nu- tions in both species are obviously not presentes en la península ibérica y norte tricias conocidas de las larvas de ambas constant in appearance in particular lo- de África en base a datos moleculares especies y se detalla la diferente colora- calities and no convincingly justification obtenidos del gen mitocondrial citocro- ción de las orugas adultas. También se can be concluded for maintaining in- ma oxidasa subunidad I (COI) y en rela- ilustran y describen las características de fraspecific distinctions from the external ción a su morfología interna y externa. los diferentes hábitats que ocupa. appearance. According to relevant diag- Se expone su distribución y rango de nostic features and genetic differentia- variabilidad. Las especies de Psilogaster Zusammenfassung. Von Populationen tion, three allopatric taxa are recognized son polimórficas en lo que respecta a su der Gattung Psilogaster Reichenbach, in Psilogaster, two treated as distinct spe- coloración y patrón alar. Esto ha supues- 1817 auf der Iberischen Halbinsel und cies, P. algeriensis (Baker, 1885) and P. to la descripción de varios sinónimos. Sin in Nordafrika wurden molekulare Da- loti (Ochsenheimer, 1810), and the lat- embargo, las variaciones en ambas espe- tensätze der mitochondrial cytochrome ter divided into two subspecies, P. loti loti cies no son aparentemente constantes en oxidase subunit I gene (COI) sowie äuße- and P. loti claussi (Huertas Dionisio, las diferentes localidades y en base a su re und innere morphologische Merkmale 1977) stat. rev., based on moderate dif- apariencia externa no se puede concluir untersucht, und ihre Verbreitung und ference in available molecular genetic que exista una justificación suficiente Variabilität werden diskutiert. Psilo data (COI mtDNA) and life history traits. para mantener las distinciones infraes- gaster-Arten sind in Bezug auf ihre Far- Psilogaster loti loti occurs throughout the pecíficas planteadas con anterioridad. De be und Zeichnung im Vorderflügel po- Iberian Peninsula north to the French acuerdo con los elementos diagnósticos lymorph. Dies gab Anlass zur Beschrei- Pyrenees, whereas P. loti claussi is lim- más relevantes y las diferencias genéticas bung einiger als konspezifisch zu wer- ited to south-western Andalusia (Huelva, observadas, se distinguen tres taxones tender Taxa, da die auftretenden habi- Sevilla and Cádiz). Psilogaster algeriensis alopátricos. P. loti loti (Ochsenheimer, tuellen Modifikationen regional nicht (Baker, 1885), confirmed as a distinct 1810) aparece desde el norte de los Piri- überzeugend eingrenzbar und deshalb species based on the high minimum ge- neos franceses y por toda la peninsula für infraspezifische Unterscheidungen netic p-distance of 3.8 % of its COI se- ibérica, desde el norte hasta las regiones nicht verwertbar sind. Die vorliegende quence from that of P. loti and on differ- más meridionales. Psilogaster loti claussi Studie belegt die Existenz von drei allo- ences in larval structures, inhabits re- (Huertas Dionisio, 1977) stat. rev., limi- patrisch verbreiteten Taxa in Psilogaster, gions in North Africa. The taxonomic tada a determinadas zonas del suroeste zwei auf Artebene, P. algeriensis (Baker, status of Diploma simulatrix Chrétien, de Andalucía (Huelva, Sevilla, Cádiz), se 1885) and P. loti (Ochsenheimer, 1810). 1910 syn. rev. is revised to that of a sub- considera una subespecie en base a su Letztere ist aufgrund moderater Ver- strate form of P. algeriensis. A neotype is biología y en vista de la existencia de tan schiedenheit in den vorliegenden mole- designed for P. loti loti, presently depos- solo una diferencia moderada con la an- kulargenetischen Daten (COI mtDNA) ited in MWM but in future in the Zoo- terior de acuerdo con los resultados ge- und ihrer Lebensweisen in zwei Unter- logical State Collection Munich. The néticos disponibles (mtDNA: COI-barco- arten, P. loti loti und P. loti claussi (Hu- known natural larval hostplants of both des). Psilogaster algeriensis (Baker, ertas Dionisio, 1977) stat. rev. unter- 85 Entomologische Zeitschrift · Schwanfeld · 128 (2) 2018 teilt. Psilogaster loti loti ist auf der Iberi- mindestens 3.8 % abweichenden mole- wahrt in MWM, später in Zoologische schen Halbinsel nördlich bis in die fran- kulargenetischen Distanz und ihrer cha- Staatssammlungen, München). Für bei- zösischen Pyrenäen verbreitet, während rakteristischen Larvalstrukturen als Art de Arten werden die in freier Natur P. loti claussi auf das südwestliche An- bestätigt. Der Status von Diplura simu nachgewiesenen larvalen Wirtspflanzen dalusien (Huelva, Sevilla and Cádiz) latrix Chrétien, 1910 syn. rev. wird re- gelistet, und die unterschiedliche Tracht beschränkt ist. Psilogaster algeriensis vidiert zu dem einer Substratform von der erwachsenen Raupen wie auch de- (Baker, 1885), die nordafrikanische P. algeriensis. Für P. loti loti wird ein ren Habitate werden beschrieben und Regionen besiedelt, wird wegen der weiblicher Neotypus designiert (aufbe- illustriert. Key words. Lepidoptera, Lasiocampidae, Pinarinae, Psilogaster, revised taxonomy, Palaearctic Region, neotype, DNA barcoding, mitochondrial DNA. dicate the existence of the three infra- scribed in Vodă, Dapporto, Dincă. & Vila Introduction specific taxa. Instead, substantial differ- (2015). LepF1 and LepR1 primers (He- Psilogaster Reichenbach, 1817 is endemic ences in larval colour pattern, hostplants bert et al. 2004) were used for the COI to the atlanto-mediterranean region and and habitat preferences suggest that three amplification and obtained gene frag- appears to be taxonomically isolated, with- taxa at different taxonomic levels are rep- ments of 658 bp maximum length. The out any obvious close relationship to other resented. However, numerous species of samples were first denatured at 92 ºC for Lasiocampidae in the Palaearctic Region. the host genera Cistus and Helianthemun 60 s, then at 92 ºC for 15 s, 48 ºC for 45 s exist in the distribution range of Psilo and 62 ºC for 150 s in five cycles and in Based on morphological differences and gaster, some of which are very variable and other 30 cycles changing the annealing genetic differentiation in the mitochon- difficult to identify, so that the exact host temperature to 52 ºC, with the final exten- drial cytochrome oxidase subunit I gene ranges of the different Psilogaster taxa are sion step at 62 ºC for seven minutes. DNA (COI), only two allopatric species and one unclear. sequences were aligned with MAFFT subspecies are recognized in Psilogaster. v7.304 (Katoh & Standley 2013). The P. loti loti (Ochsenheimer, 1810) inhabits best-fitting nucleotide substitution models the Iberian Peninsula and the French East Abbreviations according to jModelTest v2.1.7 (Darriba, Pyrenees, whereas P. loti claussi (Huertas CBB – Collection Bruer, Braunschweig; Taboada, Doallo & Posada 2012) were Dionisio, 1977), recognised as a valid sub- CDF – Collection de Freina, München, GTR+γ for COI under BIC. We performed species based on small differences in mo- CML – Collection Monasterio León, Lo- a Bayesian inference phylogeny using Mr. lecular genetic sequences and life history groño, e. l. – ex larva, DF – de Freina Bayes v3.2.6 (Ronquist et al. 2012) with traits, occurs in a small sector in south- GPdF – Genital preparation de Freina, 1000000 generations and sampling every western Spain (the coastal areas of Huel- IBEB – Institut de Biología Evolutiva 1000 generations. PCR products were pu- va, Sevilla and Cádiz). Psilogaster algerien (CSIC-UPF), Barcelona; ID – DNA identi- rified and Sanger-sequenced by Macro- sis (Baker, 1885), differing in a larger ge- fication number in MWM, München; ML gen Inc. Europe (Amsterdam, the Nether- netic divergence as well as in habitat and – Monasterio León; MWM – Museum lands). All sequences have been deposited colour pattern of adults