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 (: , 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-

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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 , 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 and last larval in- Witt, München (in ZSM); WB – Werner in GenBank. stars from P. loti, represents the genus in Bruer; ZSM Zoological State Collection the North African Maghreb region. Munich. Genetic distances between subspecies were calculated for COI using MEGA Psilogaster is a highly polymorphic genus, v7.0.14 (Kumar et al. 2016) with the boot- which has led to the description of several Methods and materials strap method to estimate variance and infraspecific taxa and weakly defined in- Morphological and phenological analysis using uncorrected p-distances (Collins dividual forms, as was fashionable in is based on numerous adults and imma- et al. 2012). Genetic distances between the last century. Their nomenclatural sta- ture stages as well as intensive observa- species are reported as minimum pair- tus was interpreted controversially in the tions of life history, both under natural wise distances, while intraspecific varia- literature (Grünberg 1913, Rougeot & conditions and in captive breeding exper- tion is reported as maximum pairwise dis- Viette 1978, de Freina & Witt 1987, iments. Genitalia were dissected, macer- tances. Leraut 2006, Lewandowski & Lewan- ated, cleaned, stained and examined in dowski-Krenz 2017). In recent years, two 30 % ethanol and the slide-mounted in Eu- DNA extracts are stored at the IBEB. All of us (ML and WB) carried out extensive paral for photography and final storage. studied specimens and samples, with iden- collecting and fieldwork on the genus in tification numbers (ID) listed below, are different regions of Spain (ML, WB) and The cytochrome c oxidase I (COI) mito- stored in the collection of MWM: Tunisia (WB). chondrial marker was analysed for seven Psilogaster specimens (Tab. 1), represent- Psilogaster loti loti: ID IDEB 16J949: The results revealed that some of the ear- ing one population from Tunisia and six Spain, Prov. Guadalajara, Cantalojas (in lier statements about the life history of distant Iberian populations. DNA extrac- coll. Aistleitner, Feldkirch); ID IDEB Psilogaster are generalized and do not in- tion was done following the protocol de- 16M092: Spain, Prov. Valencia, [Umg. Va-

86 Josef J. de Freina et al., Geographic and biological variation in Psilogaster lencia], El Saler, 1 `, ex larva, 27.I.1991, Taxonomy reads: „Gastropacha loti. Der Mann hat leg. et cult. E. M. Muńoz (CDF, in MWM); kaum die Größe von Gastr. franconia; ID IDEB 16M093: Spain, La Rioja, Ez- Psilogaster Reichenbach, 1817 seine Fühler sind rostfarbig, mit einem gel- caray. Psilogaster Reichenbach, 1817: 283. blichen Schaft, der Kopf, Rücken und Hinterleib graubraun und stark behaart, Psilogaster loti claussi: ID IDEB 16M094, Type species. Gastropacha loti Ochsenhe- die Vorderflügel von der Wurzel aus fuchs- ID IDEB 16M095 and ID IDEB 16M096: imer, 1810 (by monotypy). roth, dann rostbraun; in der Mitte steht Spain, Prov. Cádiz, Puerto Real, Fac. ein weißes rundes Fleckchen und jenseits Cienc. Educación de la Universidad de Diplura Rambur, [1866]: 350; preoccu- desselben ein weißer, etwas geschweifter Cádiz, larva 7.XII.2006, e. l. 10.III.2007, pied by Diplura Koch, 1850 (Arachnida). Streif in schiefer Richtung. Die Hinter- leg. et cult. Monasterio León, GPdF P. loti Dipluriella Strand, 1910: 14; replacement flügel nebst der Unterseite sind einfarbig claussi 2016/22 (MWM). for Diplura Rambur, [1866]. rostbraun.

Psilogaster algeriensis: ID IDEB 16J953, Psilogaster loti Das Weib ist nur um weniges größer, dur- caterpillar: Tunesien, Djerba, Trifa. chaus bräunlichgrau, mit einem weißen (Ochsenheimer, 1810) Mittelpunkte und verloschenen gleichfar- Gastropacha loti Ochsenheimer, 1810: bigen [sic] Querstreife auf den Vorder- Results 291. flügeln. Der Hinterleib hat keinen After- The results of the molecular analysis (Tab. büschel“. 1, Textfig. 1) show that the North African Type locality. Portugal. Psilogaster populations display a minimum Ochsenheimer's collection was sold to the genetic distance of 3.8 % from the Iberian The original description was based on a Hungarian National Museum in Budapest ones, inferring the existence of two dis- pair collected by Johann Centurius Graf in 1824, where it was largely destroyed by tinct species (P. algeriensis and P. loti). von Hoffmanssegg in Portugal, but a flooding in 1838. Despite intensive search- However, DNA sequence data of recently more precise locality was not provided. It es of the remaining material of the discovered closely restricted outlying pop- ulation in the Spanish southwest displays not unimportant differences to those of the rest of Spain.

As a result, P. loti comprises two subspe- cies, each one recovered as monophyletic (Textfig. 1), with not insignificant differ- ences in the mtDNA COI-barcode (1.3 % minimum distance) and, according to pre- vious knowledge, well-defined differences in biology: such as in habitat require- ments, phenology, spectrum of larval host plants and larval pattern. Both seem to ex- ist in different habitats of Huelva, Cádiz and probably Sevilla. Psilogaster algerien­ sis, which distribution is limited to North- ern Africa, markedly differs in the barcode sequences and is confirmed as a separate species. Indeed, it shows a minimum ge- netic distance to P. loti of 3.8 % in the COI (Tab. 1), as well as biological character­ Textfig. 1. COI gene tree obtained through Bayesian inference. Posterior probabilities indicated in istics. the nodes and scale units are presented in substitutions per site.

Tab. 1. Specimens used for genetic analyses and uncorrected p-distances of each individual with P. algeriensis.

Sample ID Taxon Locality Genetic distances with P. algeriensis 16J949 Psilogaster loti loti Cantalojas, Guadalajara, Spain 3.9 % 16M092 Psilogaster loti loti El Saler, Valencia, Spain 4.1 % 16M093 Psilogaster loti loti Ezcaray, La Rioja, Spain 3.8 % 16M094 Psilogaster loti claussi Cádiz, Cádiz, Spain 4.1 % 16M095 Psilogaster loti claussi Cádiz, Cádiz, Spain 4 % 16M096 Psilogaster loti claussi Cádiz, Cádiz, Spain 4 % 16J953 Psilogaster algeriensis Djerba island, Medenine, Tunisia – KX045895 lobulina – 8.5 %

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1 2 3 4

5 6 7 8

9 10 11 12

13 14 15 16

17 18 19 20

21 22 23 24 25

26 27 28 29 30

88 Josef J. de Freina et al., Geographic and biological variation in Psilogaster

Ochsenheimer collection, no other syn- whether this effect is due to ecological fac- cocoons are elongate ovoid, pale silvery types could be found (László Ronkay, tors during the pupal stage or a genetical- gray-brown in colour and densely woven Hungarian Natural History Museum, Bu- ly coded polymorphism. Breeding experi- with a hard outer layer, interspersed with dapest, personal communication). In view ments show that the coloration is largely black bristles. The larvae are heavily para- of the apparent lack of any types remain- temperature-dependent. The minimum sitized (up to 80 %; Pérez de Gregorio ing, and to stabilise the nomenclature of difference in COI between this subspecies 1976) by tachinid flies such as Exorista seg­ the species, we here designate a female and P. loti claussi was found to be 1.3 %. regata and wasps such as Chalcididae. specimen agreeing with Ochsenheimer's description as a neotype (acc. to Article 75 Habitats, life history and hostplants. Distribution. Psilogaster loti loti occurs in of the ICZN (1999), with the following de- Psilogaster loti loti inhabits a large variety most parts of the Peninsula but never in tails: Portugal, Prov. Alentejo, Lagoa de of habitats in which its hostplants occur. abundance. It has been recorded from Ma- Santo André, 3 km W Vila Nova, 5 m, It is usually found in rocky semi-humid ar- drid, Teruel, Zaragoza, Huesca, León, Bur- 25.IX.2009, leg. P. Skou. coll. CDF (Fig. eas dominated by Mediterranean vegeta- gos, Salamanca, Soria, Valladolid, Zamo- 2). The definition of a neotype is necessary tion, such as Quercus ilex, Pinus spp., Ros­ ra, Palencia, Sevilla, Granada, Málaga, to distinguish the taxon loti from the ­other marinus officinalis or Buxus sempervirens, Cádiz, Huelva, Navarra, La Rioja, Álava- nominal Psilogaster taxa. The neotype is and occurs over a large altitudinal range, Araba, Girona, Barcelona, Lleida, Tarra- currently deposited in MWM but will later from sea level to at least 1500 m, and pre- gona, Cuenca, Orense, Lugo, Pontevedra, be housed in ZSM. sumably higher in northern Spain. Al- Cáceres, Badajoz, Murcia (Barco & San- though considerable climatic differences tamaria-Hernandez 2011; Blat Beltrán Psilogaster loti loti exist between the northern and southern 1976; Bolland 1976; Cifuentes, Borruel areas of the Iberian Peninsula where the & Plaza 1993; De Ibarra 1980a, 1980b; (Ochsenheimer, 1810) taxon occurs, it is bivoltine throughout its Derra & Hacker 1982; Hermosa & Gómez (Figs 1–9, 21–23, 31–34, 41–44, 50–53) range. Depending on altitude, the first- Bustillo 1974; Huertas Dionisio 1979; Diplura loti var. vernetensis Oberthür, generation adults emerge in April, May Magro & Jambrina 2013; Méndez 1983; 1916: 326–327. and June and the second from July to Moreno Tamurejo 2002; Murria Belt- ­October, and their respective larvae can rán 2012; Novoa Pérez, Nieto Manzano, Type locality. Vernet-les.Bains, French East be found from May to August and from GarcÍa-Villanueva & Morena Tamurejo Pyrenees. August to October (Huertas Dionisio 2002; OrtÍz, de la Calle, Rubio, Garre 1979). The larvae feed on a large number & Guerrero 2007; Pérez De-Gregorio Variability/Remarks. Psilogaster loti loti is of Cistaceae, such as Cistus salvifolius, 1976, 1977; Pérez de Gregorio, Maso very variable, but there is no considerable C. albidus, C. populifolius, C. ladanifer, Planas & Castells 1977; Redondo 1976, clear geographical variation. The main C. laurifolius, C. clusii, C. monspeliensis, 1986; Ribbe 1909-1912; Vaamonde & Es- characters used by authors to distinguish Halimium umbellatum and tévez Rodríguez 1991; Vallhonrat, infraspecific taxa are differences in ground apenninum (Murria Beltràn personal ob- Cervello, Marti, Pérez de Gregorio & colour, which may vary from greyish servation, for Huesca and Aragon). Gómez Xaus 2005; Vega Escandón 1977, 1979 brown to chocolate-brown. Additional dif- Bustillo & Fernández-Rubio (1976) also and personal observations). On the Inter- ferences also occur in the more or less pro- suggested Rosmarinus officinalis (Lami- net there are also photos of specimens nounced white antemedial transverse line aceae) as a larval host, but we doubt this from Castellón, Segovia, Ciudad Real, and the white discal spot in the forewing, to be a natural foodplant. Preferred larval Murcia, Alicante, and Almería. The only which can be absent. habitats are sparsely vegetated and in full Spanish areas from which it has not been sun or partially shaded. Larval coloration reported are Asturias and Cantabria. These and some variation in size can be varies moderately and mainly affects the found in any population, but mainly in proportions of red and yellow and the Psilogaster loti claussi southern Spain. Specimens in lower and prominence of the white spots (Figs 31– warmer regions are typically paler and 34). Most larvae pupate before winter, (Huertas Dionisio, 1977) those in higher areas with more humid spinning a cocoon close to the ground be- (Figs 10–15, 24, 25, 35–38, 45, 46, 53–55) ­climate and lower temperatures darker tween leaves or among litter and usually Dipluriella loti ab. homochroa Zerny, (von Buddenbrock 1961). It is unclear some distance from their hostplant. The 1927: 64; infrasubspecific.

Figs 1–9. Psilogaster loti loti. – 1. `, Portugal, Setubal, ex larva 10.IX.80 [larva M.V.80], [leg. et cult.] R. Menrad, coll. De Freina (MWM). 2. ´, neotype, Portugal, Prov. Alentejo, Lagoa de Santo André, 3 km W Vila Nova, 5 m, 25.IX.2009, P. Skou leg. 3. `, Spain, Prov. Girona, Puig de Ventós, Vidreres, 60 m, e. l. 26.VI.1990, A. E. Rau leg. 4. ´, Spain, Barcelona, e. l. 22.V.1922, W. Marten leg. (illustrated by de Freina & Witt 1987, pl. 26, fig. 47). 5. `, [Spain], Aragón, Albarracín, e. l. 5.VIII.1927, [leg.] Predota (illustrated by de Freina & Witt 1987, pl. 26, fig. 44). 6. `, [Spain], Aragón, Albarracín, e. l. 8.VI.1933, A. Schleppnik leg. 7. `, South Portugal, e. l. 7.IV.1933, W. Marten leg. 8. ´, South Portugal, e. l. W. Marten. 9. ´, Southern Spain, [Prov. Granada], Guadix. – Figs 10–15. Psilogaster loti claussi. – 10. `, Spain, Huelva, El Rocio, e. l. 31.V.1938, W. Marten leg. (illustrated by de Freina & Witt 1987, pl. 26, fig. 48). 11. `, Spain, area Huelva, Punta Umbria, e. p. 15.X.2006, Antonietty & Monasterio León leg. et cult. 12–14. ´, e. l. (Figs 12, 13 forma homochroa), [Spain, Cádiz], Puerto Real, Fac. Cienc. Educación de la Universidad de Cádiz, larva 7.XII.2006, e. l. 10.III.2077, Monasterio León leg. et cult., GPdF 2016/23 (MWM). 15. `, e. l., same data as Fig. 11 (GPdF 2016/22 (MWM). – Figs 16–20. Psilogaster algeriensis. – 16. `, Algeria, Predota. 17. `, Algeria, Biskra, 1.VIII.1933 (illustrated by de Freina & Witt 1987, pl. 26, fig. 46). 18. ´, [Tunisia], Kahiri c., Gafsa, 14.VIII.1909 (Helianth.[emum]). 19, 20. `, ´, ­Tunisia, Djerba, Trifa, 5–10 m, M.IV.2016, Bruer leg. et cult. (specimens 1–18 in MWM, 19–20 CBB). – Figs 21–23. Psilogaster loti loti. – 21, 22. `, ´, e. l., ` hatched 12.IV.2008, ´ with cocoon, hatched 21.III.2009, N. Spain, Prov. Huesca, Aineto, Murria Beltrán cult. 23. `, Ezcaray (Prov. La Rioja), 16.IV.2014, Monasterio León leg. – Figs 24, 25. Psilogaster loti claussi. – 24. Hatched `, Andalusien, Umg. Mazagón, ca. 40 m, Bruer cult. 25. Hatched ´ with co- coon, same data as Fig. 12. – Figs 26–30. Psilogaster algeriensis, same data as Fig. 19. 26. Cocoon. 27, 28. Hatched `. 29, 30. Hatched ´. – Photos­ 21, 22 Murria Beltrán, 23, 25 Monasterio León, 24, 26–30 Bruer.

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31 32

33 34

35 36

37 38

39 40

90 Josef J. de Freina et al., Geographic and biological variation in Psilogaster

Dipluriella loti claussi Huertas Dionisio, er. This subspecies is also bivoltine. The loti are not clear, and it is unknown wheth- 1977: 64. first-generation adults occur from March er a transition zone may occur between to June and those of the second from June them. Type locality. Zona colindante a Doñana, to September, and their respective larvae Almonte, Provinz Huelva. from April to August (most commonly at Psilogaster algeriensis Dipluriella loti f. contrariata Huertas Di- the end of April) and from late summer to onisio, 1977: 64; infrasubspecific. March (at Donana and El Rocina; Bruer, (Baker, 1885) Dipluriella loti f. nigrolineata Huertas Di- personal observations). Sometimes adults (Figs 16–20, 26–30, 39, 40, 47–49) onisio, 1977: 64; infrasubspecific. or larvae of the two generations occur con- Bombyx loti var. algeriensis Baker, 1885: Dipluriella loti f. nigromaculata Huertas temporaneously and a third generation is 242. Dionisio, 1977; 61; infrasubspecific. faked, when the last second-generation Dipluriella loti f. serrdorensis Huertas Di- adults fly in September and larvae hatched Type locality. Algeria, Guelfa. onisio, 1977; 61; infrasubspecific. in July have already spun their cocoons. Diplura loti brunnea f. brunnea Oberthür, Some pupae of the first generation also 1900: 338; infrasubspecific. Variability. Compared with the nominate aestivate and only hatch a year later Diplura simulatrix Chrétien, 1910: 78 subspecies, P. loti claussi is more uniform (Huertas Dionisio 1979). The larvae feed Type locality. Tunesien, Gafsa. in habitus and varies only slightly in pat- on Halimium halimifolium, Cistus salviifo­ tern. It differs mainly by its darker colora- lius, C. monspeliensis (Huertas Dionisio Variability. Psilogaster algeriensis is a well- tion, culminating in the form homochroa, 1979, 2007), , Cistus populi­ distinct species, both morphologically and in which the white antemedial transverse folius, , , genetically, the difference in its COI se- line and the white discal spot on the fore- . The reference of Quercus as quence from that of P. loti amounting to wings are absent. The subspecies has a ten- hostplant (Aue 1933) is inaccurate, as is at least 3.8 %. The adults are entirely pal- dency to develop monochromatic dark fe- that of Rosmarinus officinalis. The mature er yellowish to pale brownish or ochreous- males. It also differs in the colour of its larvae, which feed exposed and singly, dif- brown in colour. Tunisian specimens are larva, hostplant preferences and lifecycle, fer from those of P. loti loti in their con- sometimes smaller, the whitish zig-zac and the specimens analysed genetically spicuous orange-yellow bristles and larger markings more pronounced than in typical display a 1.3 % minimum difference in white spots (especially posteriorly and lat- P. algeriensis. These represent the form their COI sequences from P. loti loti. How- eroventrally) and the absence of red spots simulatrix Chrétien, which, however, ever, no genital differences are detectable laterally (Figs 37, 38; Figs 9–11 in Lewan- does not occur constantly and is not pre- between the subspecies (Figs 50–55); P. dowski & Lewandowski-Krenz 2017). dominant, and there is no justification ev- loti loti. ´, GPdF 2016/18; ´, GPdF 2016/19 The larvae are also frequently parasitized ident to recognize it as a subspecies. The (Aragón; Prov. Huesca, Barranco de Ur­ by Tachinidae, the rate in some years differences in the female genitalia be- gellet, Villanueva de Sigena, 250 m, reaching almost 50 % (pers. obs. WB). tween P. algeriensis and P. loti as detailed 30TYM4721, 22.VIII.1997, leg. R. Macià, by Leraut (2006) were not checked by us. coll. de Freina, MWM); `, GPdF 2016/20, Distribution. Psilogaster loti claussi only The larva differs conspicuously from that ´, GPdF 2016/21 (Prov. Valencia, [Umg. occurs in a small coastal area of the prov- of P. loti, especially in the last two instars. Valencia], El Saler, 1 m, ex larva, 27.I.1991, inces Huelva, Cádiz and Sevilla, near Do- It lacks the white dorsolateral patches and leg. et cult. E. M. Muńoz, coll. de Freina, ñana. The best-known populations are the dorsal setal tufts are orange and the MWM); P. loti claussi. `, GPdF 2016/22, around Lepe-Cartaya, Punta Umbría-El dorsolateral ones vermilion red. It is sig- ´, GPdF 2016/23 (Prov. Cádiz, Universi- Rompido, Almonte, Palos de la Frontera, nificantly larger, and the female larva is dad de Cádiz, larva 7.XII.2006, e. l. 10. Lucena del Puerto, El Rompido, Barbate, conspicuously larger than the male. A de- III.2077, leg. et cult. Monasterio León, Mazagón, Matalascañas (Huelva) and tailed description of all larval instars, coll. de Freina, MWM). Puerto Real (Cádiz). The taxon is present which agrees with our observations, is in almost every protected area along the ­given by Lewandowski & Lewandowski- Habitats, life history and hostplants. Huelva coast: Marismas del Río Piedras y Krenz (2017, figs 1–8). Psilogaster loti claussii inhabits sandy Flecha del Rompido Natural Park, Laguna coastal biotopes with a vegetation com- del Portil Natural Reserve and Protection Habitats, life history and hostplants. posed mainly of Pinus, Quercus ilex and Area, Enebrales de Punta Umbría Natural Psilogaster algeriensis is known to occur in shrubs such as Pistacea, Aristolochia bae­ Park, Marismas del Odiel Natural Park, Es- low to colline landscapes, in halophilic tica, Halimium halimifolium and Cistus tero Domingo Rubio Natural Park, Lagu- and very sparsely vegetated habitats, with spp., always in low altitudes from 0 to nas de Palos y las Madres Natural Park temperatures similar to those of south- 15 m above sea level. Winter temperatures (Huertas Dionisio 2007), as well as in western Andalusia but less precipitation. (in December) are similar to those in the Doñana National Park. The borderline Mature larvae are found only from Novem- North Africa, but the annual rainfall is low- between its populations and those of P. loti ber until the end of January. At Djerba,

Figs 31–40. Preimaginal adult instars (L5). – 31–34. Psilogaster loti loti. – 31. Spain, Prov. Huesca, Aineto, lingering casually at the non hostplant Genista scorpius. 32. Spain, Prov. Huesca, 10–20 km S Candasnos, Barranco de la Valcuerna, 400 m, 18.III.2007. 33, 34. Spain, Limit between Provs. Araba-Álava and La Rioja. – Figs 35–38. Psilogaster loti claussi. – 35, 36. Spain, Huelva, Mazagón, 15.IV.2015, feeding on Halimium halimifolium. 37, 38. [Spain, Cádiz], Puerto Real, Fac. Cienc. Educación de la Universidad de Cádiz, 7.XII.2006, feeding on Halimium halimifolium. – Figs 39, 40. Psilogaster algeriensis, same data as Fig. 19, feeding on Helianthemum confertum, Fig. 39 dorsolateral, Fig. 40 lateral, dorsal. ´. – Photos 31 Murria Beltrán, 32 de Freina, 33, 34, 37, 38 Monasterio León, 35, 36, 39, 40 Bruer.

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41 42 43

44 45 46

47 48 49

51

50

52

55

53 54

92 Josef J. de Freina et al., Geographic and biological variation in Psilogaster south of Trifa, we (WB) encountered them material available for DNA analysis. Thaumetopoeidae, Thyretidae, Axiidae, Dre­ in a large area five to ten metre above sea Harald Sulak, MWM, assisted with pho- panidae,­ Thytiridae, Bombycidae, Brahmae­ level from the middle of December until tography and Tibor Csövari, Budapest, idae, Endromidae, Lasiocampidae, Lemoni­ idae, Saturniidae, Sphingidae). 708 S., 47 the first week of January, pupating in prepared genitalia. Wolfgang Nässig, Farbtafeln. Edition Forschung & Wissenschaft, ­captivity from the end of December until Forschungsinstitut Senckenberg, Frank- München the middle of January. The pupal stage furt am Main, very kindly assisted with ap- De Ibarra, M. 1980a. Mis memorias (Cuarta par- lasted more than twelve weeks, the ima- plication of the rules of zoological nomen- te). SHILAP, Revista de Lepidopterología 9 (35): gines only hatching between in late April. clature (ICZN), and Rolf Oberprieler, 233–237. De Ibarra, M. 1980b. Mis memorias (Cuarta par- Lewandowski & Lewandowski-Krenz CSIRO, Canberra, is thanked for linguistic te). SHILAP, Revista de Lepidopterología 9 (36): (2017) reported a pupal stage lasting be- improvements of the manuscript. Finally 303–307. tween five and twelve weeks. The larvae we wish to thank Eva Karl, ZSM, who has Derra & Hacker H. 1982. Contribution to the prefer habitats in full sun or half shade. been very helpful in tracing literature. lepidóptera-fauna of Spain. Heterocera of a The only recorded host plant for the pop- three-week visit in summer 1980 (x) (II). ulation at Djerba is Helianthemum confer­ SHILAP, Revista de Lepidopterología 10 (37): References 23–31. tum (WB, pers. obs.), but for the Maghreb Gómez Bustillo, M. R. & Fernández-Rubio, F. numerous Helianthemum spp. have been Aue, A. U. E. 1933. Handbuch für den praktischen 1976. Mariposas de la Península Ibérica, Heteró­ reported, such as H. ellipticum, H. erioce­ Entomologen. Band 4, 180 pp. [D. loti p. 33]. ceros I Vol III. 300 pp. Servicio de Publicaciones phalum, H. geniorum, H. hirtum. As with Verlag des Internationalen Entomologischen del Ministerio de Agricultura, Madrid. P. loti, the larvae are also parasitized by Vereins e. V., Frankfurt am Main. Grünberg, M. 1933. Die Groß-Schmetterlinge der Baker, G. T. 1885. Descriptions of some new spe- Tachinidae. Erde. Eine systematische Bearbeitung der bis cies of Lepidoptera from Algeria. The Entomo­­ jetzt bekannten Groß-Schmetterlinge. In Seitz, logist's Monthly Magazine 21 (4): 241–245. A. (Hrsg.) Band 2, Die palaearktischen Spinner Distribution. The species is restricted to Blat BeltrÁn, F. 1976. Cazaderos de mariposas und Schwärmer, vii + 315 S., 16 Taf. [Lasiocam- North Africa. The incorrect reference for en los Montes Universales y sierra de Albarracín pidae S. 109–125, Taf. 9, 10]. A. Kernen, Stutt- southern Spain (Leraut 2006) has already (VII) (Resumen de doce temporadas de caza) gart. been corrected by Lewandowski & Lewan- (Continuación). SHILAP, Revista de Lepidopte­ Hebert, P. D. N., Penton, E. H., Burns, J. M., rología 15: 237–242. Janzen. D. H. & Hallwachs, W. 2004. Ten spe- dowski-Krenz (2017). Psilogaster alge­ Bolland, F. 1976. Province de Huesca: Paradis cies in one: DNA barcoding reveals cryptic di- riensis appears to be a scarce species in the entomologique. SHILAP, Revista de Lepidopte­ versity in the neotropical skipper butterfly As­ Maghreb, where it is confined to the costal rología 14: 135–142. traptes fulgerator. PNAS 101: 14812–14817. and colline areas in Tunisia and Algeria, Buddenbrock, W. von 1961. Einige Bemerkungen Hermosa, E. & Gómez-Bustillo, M. R . 1974. Poe­ as far as known. However, no proper über die Biologie der andalusischen Rasse von cilocampa canensis (Mill., 1877), nueva especie searches for it in the region have been car- Diplura loti O. Nachrichtenblatt der Bayerischen para la Península Ibérica (Lep. Lasiocampidae). Entomologen 10 (8): 81–85. SHILAP, Revista de Lepidopterología 5: 46–49. ried out during the winter months. Chrétien, P. 1910. Description de nouvelles es- Huertas Dionisio, M. 1979. Dipluriella loti (Ochs- pèces de Lépidoptères de Tunisie. Le Natura­ enheimer, 1810) en Huelva y descripción de Acknowledgements. We are greatly in- liste, Revue illustrée des Sciences Naturelles (2. una subsp. y cuatro formas nuevas (Lep. Lasio- debted to Roger Vila, Institut de Biologia Sér., 22), 32: 78–79. campidae). SHILAP, Revista de Lepidopterologia Cifuentes, J., Borruel, M. & Plaza, B. 1993. 7 (25): 59–68. Evolutiva (CSICUPF), Barcelona, who Catálogo y atlas de los lepidópteros macroheteró­ Huertas Dionisio, M. 2007. Lepidópteros de los generously made his laboratory available ceros de Navarra. 235 pp. Gobierno de Navarra, Espacios Naturales Protegidos del Litoral de for the molecular analyses. Ruth Escobés Pamplona. Huelva (Macro y Microlepidoptera). Sociedad Jiménez (La Rioja) helped with fieldwork. Collins, R. A., Boykin, L. M., Cruickshank, R. H. Andaluza de Entomología. Monográfico 2: László Ronkay, Hungarian Natural His- & Armstrong, C. F. 2012. Barcoding's next top 1–250. model: an evaluation of nucleotide substitution tory Museum, Budapest, kindly searched Katoh, K. & Standley, D. M. 2013. MAFFT mul- models for specimen identification. Methods in tiple sequence alignment software version 7: for type material in the Ochsenheimer Ecology and Evolution 3: 457–465. improvements in performance and usability. collection. Enrique Murria Beltrán, Darriba, D., Taboada, G. L., Doallo, R. & Posada, Molecular Biology and Evolution 30: 772–780. Aineto (Huesca) and Arturo Baquero, D. 2012. jModelTest 2: more models, new heu- Kumar, S, Stecher, G. & Tamura, K. 2016. Sevilla, provided information, photos and ristics and parallel computing. Nature Methods MEGA7: Molecular Evolutionary Genetics Anal- fieldwork. Carlos Antonietty, Sevilla, 9: 772–772. ysis Version 7.0 for Bigger Datasets. Molecular De Freina, J. J. & Witt, T. J. 1987. Die Bombyces Biology and Evolution (7): 1870–1874. E. M. Muńoz, Valencia, R. Macià, Barce- 33 und Sphinges der Westpalaearktis (Insecta, Lepi­ Leraut, P. 2006. Lasiocampidae. S. 66–87. In: lona, Eyjolf Aistleitner, Feldkirch, and doptera) Vol. 1 (Nolidae, Arctiidae, Syntomi- of Europe, Vol. 1. 396 pp. N. A. P. Editions, ­Stefan Lewandowski, München made dae, Dilobidae, Lymantriidae, Notodontidae, Verrières le Buisson.

Figs 41–44. Habitats of Psilogaster loti loti. – 41. Spain, Sierra de Guara, vic. Lusera (Huesca), 800 m, local larva on Helianthemum apenninum (Photo 19.VI.2015). 42. Spain, 15 km SW of Candasnos (Huesca), Los Monegros, 350 m, local larva on Cistus clusii (Photo 18.III.2007). 43. Spain, near Sepúlveda (Segovia), Villar de Sobrepeña, 1000 m, local larva on Cistus sp. (Photo 21.VI.2013). 44. Spain, Puerto de Pozazal S Reinosa (Cantabria), 1200 m, local larva on Helianthemum sp. (Photo 22.VII.2010). – Figs 45, 46. Habitat of Psilogaster loti claussi, Nationalpark Dońana, Huelva (Photo 21.IV.2015). – Figs 47–49. Habitat of Psilogaster algeriensis, Tunisia, Djerba, Trifa, 5–10 m. 49. Hostplant Helianthemum confertum with adult instar (insertion) (Photo M.VI.2015). Photos 41–44 de Freina, 45–49 Bruer. – Figs 50–55. Ptiologaster loti, genitalia ventral, phallus lateral (a) and pattern of andominal tergites/- sternites, lateral (b). – 50. `, Psilogaster loti loti, GPdF 2016/18, Aragon, Prov. Huesca, Barranco de Urgellet, Villanueva de Sigena, 250 m, 30TYM4721, 22.VIII.1997, R. Macia leg. 51. ´, Psilogaster loti loti, GPdF 2016/19, data as Fig. 50. 52. `, Psilogaster loti loti, GPdF 2016/20, Prov. Valencia, [Umg. Valencia], El Saler, 1 m, ex oruga 27.I.1991, E. M. Muńoz leg. et cult. 53. ´, Psilogaster loti loti, GPdF 2016/21, data as Fig. 52, but ex oruga 2.V.1993. 54. ´, Psilogaster loti claussi, GPdF 2016/23, [Spanien], Prov. Cádiz, Puerto Real, Fac. Cienc. Educación de la Universidad de Cádiz, larva 7.XII.2006, e. l. 10.III.2077, Monaster- io León leg. et cult. 55. `, Psilogaster loti claussi, GPdF 2016/22, data as Fig. 54 (all GPdF coll de Freina, MWM). Scales 5 mm. Photos 41–44 de Freina, 45, 46 Monasterio León, 47–49.

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