Molecular systematics and morphological

Zootaxa 3915 (1): 001–051 www.mapress.com /zootaxa / Copyright © 2015 Magnolia Press

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http://dx.doi.org/10.11646/zootaxa.3915.1.1 http://zoobank.org/urn:lsid:zoobank.org:pub:C23FCF79-6C86-4630-AB65-15DBEE9D51E3

Molecular systematics and morphological identification of the cryptic species of the genus Acalles Schoenherr, 1825, with descriptions of new species (Coleoptera: Curculionidae: Cryptorhynchinae) ANDRÉ SCHÜTTE1 & PETER E. STÜBEN2 1

ZFMK: Zoologisches Forschungsmuseum Alexander Koenig, Molecular Taxonomy & Biobank, Adenauerallee 160, D-53113 Bonn, Germany. E-mail: [email protected] 2 Curculio Institute, Hauweg 62, D-41066 Mönchengladbach, Germany. E-mail: [email protected]

Table of contents Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Catalogue of Acalles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Discussion and results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Acalles tree . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Acalles echinatus species complex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Acalles maraoensis species complex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Acalles sierrae species complex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Are keys for the identification of insect species still up to date? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Key to the species of Acalles Schoenherr, 1825 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Family: Curculionidae Latreille, 1802 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Subfamily: Cryptorhynchinae Schoenherr, 1825 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Genus: Acalles Schoenherr, 1825, type species: Curculio camelus Fabricius, 1792 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 New species of the subgenus Acalles s. str. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Acalles iblanensis Stüben sp. n. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Acalles vorsti Stüben sp. n. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 New species of the subgenus Origoacalles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 Acalles granulimaculosus Stüben sp. n. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 Excursus: The New World species of "Acalles" . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

Abstract Molecular systematics and morphological study of the monophyletic weevil genus Acalles Schoenherr, 1825 are presented. Based on the mitochondrial CO1 barcoding gene and 16S ribosomal RNA gene, we discuss three difficult species complexes in the framework of a molecular phylogenetic reconstruction of 37 of 47 Western Palaearctic Acalles species or subspecies: the A. echinatus, A. maraoensis and A. sierrae complexes. Two results are given: 1. An exclusive focus on morphological, exoskeletal methods reach their limits in the case of many cryptic Cryptorhynchinae. In these cases molecular analysis is indispensable to resolve species level questions. 2. By using a combination of phenotypic and genotypic characters it is not only possible to ascertain phylogenetic relationships, but also to uncover new morphological, non-intraspecifical characteristics. Digital photography with image stacking makes this possible: for the first time we present photo key for Acalles species, a reliable, less costly and quick method for identification alongside DNA barcoding. The following taxonomic changes are given: Coloracalles edoughensis Desbrochers, 1892 comb. nov. (formerly Acalles edoughensis) from North Africa and Spain change to Coloracalles Astrin & Stüben, 2008 and Pseudodichromacalles xeAccepted by R. Anderson: 3 Dec. 2014; published: 2 Feb. 2015

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rampelinus Wollaston, 1864 comb. nov. from the Canarian Island Tenerife, Acalles bazaensis Stüben, 2001 syn. nov. is a junior synonym of Acalles sierrae H. Brisout, 1865. Two new species of Acalles s. str. , A. iblanensis Stüben sp. nov. from Morocco and A. vorsti Stüben sp. nov. from Spain (Mallorca), and a new species of the subgenus Origoacalles Stüben & Astrin 2010, A. granulimaculosus Stüben sp. nov. from La Gomera, are described. Acalles temperei Péricart, 1987 stat. nov. is a subspecies of A. parvulus Boheman, 1837. A catalogue of all 43 (+4 incertae sedis) species of Acalles is presented. Finally and for the first time we compare 9 of 12 known North American so-called “Acalles” species with the Western Palaearctic species of Acalles surrounding the type species Curculio camelus Fabricius, 1792. The morphological and molecular analysis for the New World Acalles show that none of the species from the United States actually belong to the genus Acalles or one of the other genera of Western Palaearctic Cryptorhynchinae. There is one exception: Acalles costifer Le Conte, 1884, is transferred to the phylogenetically basal genus Acallocrates Reitter, 1913 as Acallocrates costifer (LeConte, 1884) comb. nov. Key words: Acalles, Origoacalles, Bayesian analysis, Integrative Taxonomy, Morphology, CO1, 16S, new species, new combination, Western Palaearctic, Spain, Canary Islands, North America, Mexico

Introduction At the end of the last century, just 150 species of the cryptorhynchine genera Acalles and Echinodera were known within the Western Palaearctic. When we subjected them to a first revision (Stüben & Behne 1998, Stüben 1998, 1999a, b), it became evident very quickly that a super-genus "Acalles" could not be retained in a phylogenetic systematics approach. The last major revision was almost 100 years ago. A. & F. Solari (1907) started a division into groups more as a quick-fix than actually referring to strict morphological or ecological criteria. This division would not last for long because more and more ecological niches inhabited by cryptic Cryptorhynchinae have been discovered since the beginning of the 21st century inhabited by more species of wingless and nocturnal Cryptorhynchinae. Explicit forest species developing in dead wood (as it is unfortunately still claimed today) are just a few, namely the southern European Echinodera species and the central European Acalles and Kyklioacalles species as well as Echinodera hypocrita from this same region. Contrary to what has been stated previously (Rheinheimer 2010), the biology of the currently known Acalles s. str. and its related genera (e.g. Onyxacalles or Kyklioacalles) is highly diverse. The latter feed on annual and perennial lignified plants, like Fabaceae, Asteraceae, Apiaceae or even succulent Euphorbiaceae and Crassulaceae. For example, the association with the Crassulaceae genus Aeonium (tree houseleek) has been used as a basis to name the genus Aeoniacalles Stüben & Astrin, 2010. Hardly a plant family is shunned by Cryptorhynchinae. Based on the discovery of new ecological niches and the co-evolution (parallel cladogenesis) of the weevils and their host plants on the Macaronesian Islands (Stüben 2000a, b; Stüben & Astrin 2010a), many new species have been described. Today nearly 400 Cryptorhynchinae species have been discovered in the well-researched Western Palaearctic. With the discovery of additional species, morphological research led to a reevaluation of structural diversity within Acalles. The most important finding here was the diversity in shape of the internal sac structure of the aedeagus. By using the endophallus, a genus or group assignment was easily possible for many Cryptorhynchinae and can often be used as a 'fingerprint' (Stüben 1998, 1999a, Stüben 2000c, Stüben & Schütte 2013a). These assignments are consistent with the ecological associations mentioned in the paragraph above. With these findings it was possible to break up the super-genus "Acalles"—in which, it seems, most authors placed everything that feigned death for longer than just a few seconds (thanatosis)—into a set of well-defined genera consistent with morphology and ecological habits. The genus Acalles has been confirmed by molecular analysis in the last few years. A broad change like this would not have been possible only by using the exoskeletal characters. Facing the inability to fly, it is futile to imagine species of this genus (apart from invasive species) being found in America, Hawaii, Australia or New Zealand as is sometimes claimed (Rheinheimer 2007, 2008). The previously known distribution of the few Western Palaearctic flightless Acalles species was from the Urals to the Caucasus and the Middle East to western Europe and northwest Africa. Nowadays 47 species are known, half of them from western Europe (France, Spain). A significant number can be sifted from leaf and twig litter, especially in old forest sites with a long forest origin (Buse 2012). They can also be caught during the spring months in Central Europe by the beating branches and twigs. Development does not take place in deadwood

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exclusively. Oviposition is done in fresh fractures of plant or tree-segments (windbreak or rockfall). The legless endophytic larvae do not produce long caverns, behaving passively in the nutrient-rich cambium and feeding on the phloem. The 12 species of the Acalles sierrae group are located in the far southwest of Europe (Iberian Peninsula) and northwest Africa (Morocco, fig. 7). It is interesting to note that they mainly occur on Fabaceae (Stüben 2004). All these species are montane open land species that have been sifted only in a few cases in detritus from scattered and shady Quercus stocks, but if present then always in combination with Fabaceae near the trunk. However, we believe the Fabaceae as host plants only play a secondary (adaptive) role, like for many other Cryptorhynchinae. The long-lasting isolation between various mountain ranges and the high altitude, mostly above 1300 m, is the main reason for the great diversity of species within this Acalles-group. Until now, the authors have had little knowledge about Cryptorhynchinae which are related to the Western Palaearctic Acalles s.l. genera (see Tree 1 and Excursus: The New World species of "Acalles"). On cursory inspection, Cryptorhynchinae species from the New World (we have added some to the analysis—see specimens marked with "New World Cryptorhynchinae" in Tree 1) look very similar to the Western Palaearctic Acalles, Onyxacalles, Kyklioacalles or Dichromacalles species, but after morphological examination of the aedeagus (endophallus) and also by molecular analysis they always turned out to be distantly related to the Western Palaearctic ones. Obviously they have undergone convergent evolution in the tropics and subtropics (see Excursus: The New World species of "Acalles"). Issues and objectives of this manuscript. In this study of the genus Acalles we examine the following three questions: 1. How helpful is the molecular analysis of the mitochondrial genes CO1 and 16S for phylogenetic reconstruction of the genus Acalles? 2. What can be accomplished by DNA barcoding (CO1) in individual cases of morphologically hard to separate (cryptic) Acalles species? 3. Aside from DNA barcoding, is there a reliable, practical and fast method of type (re-) identification by illustrations (pictoral keys)? The first two objectives we will discuss in part 4 "Results and discussion", taking three challenging species complexes into account. Here we will focus on the third question. The answer will provide an important contribution to applied entomology in the context of 'integrative taxonomy' (part 5). In part 6 'Taxonomy' we introduce some new species of the genus Acalles, and reveal new insights into the relationships of the New World "Acalles" and how these relate to the Acalles of the Palearctic Region.

Catalogue of Acalles Species included in the molecular analysis are printed in bold. l.t. = country or island containing the type locality Genus: Acalles Schoenherr, 1825; type species Curculio camelus Fabricius, 1792 Subgenus: Acalles s. str. alcarazensis Stüben, 2009; Spain almeriaensis Stüben, 2001; Spain asniensis Stüben, 2003; Morocco biokovoensis Stüben, 2008; Croatia breiti A. & F. Solari, 1909; Spain (Mallorca) camelus Fabricius, 1792 (Curculio); Austria, Belgium, Bosnia-Herzegovina, Bulgaria, Croatia, Czech Rep., Denmark, Estonia, France, Germany (l.t.), Hungary, Italy, Latvia, Lithuania, Luxembourg, Poland, Romania, Serbia, Slovakia, Slovenia, Sweden, Switzerland = nodulosis Piller & Mitterpacher, 1783 (Curculio) [nomen oblitum]

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TREE 1. (upper part): Bayesian consensus tree (50% majority rule) for CO1 and 16S, numbers at branches indicate Bayesian posterior probability (BPP) values, lower values than 100 have been deleted for better readability. New World Cryptorhynchinae contain specimens from Canada, Mexico and USA. Species names are followed by Collector´s no and ZFMK-Biobank number, former species names are provided in brackets .


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TREE 1 (middle part)



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TREE 1 (lower part)


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= quercus Boheman, 1844 = sophiae Tschapek, 1873 = variolosus Stierlin, 1887 cazorlaensis Stüben, 2004; Spain cytisi Stüben, 2004; Spain dieckmanni Péricart, 1989; France (Corsica, l.t.), Italy (Elba) dubius A. & F. Solari, 1907; Austria, France, Germany, Italy (l.t.), Slovenia, Switzerland echinatus Germar, 1824 (Cryptorhynchus); Albania, Austria (l.t.), Bosnia-Herzegovina, Bulgaria, Croatia, Czech Rep., Finland, France, Germany, Hungary, Italy, Montenegro, Poland, Romania, Russia, Serbia Slovakia, Slovenia, Sweden, Switzerland, Ukraine = echinatus var. squamosus A. & F. Solari, 1907 fallax Boheman, 1844; Bosnia-Herzegovina, Croatia, Czech Rep., France (l.t.), Germany, Greece, Hungary, Italy, Macedonia, Montenegro, Poland, Romania, Serbia, Slovakia, Slovenia, Spain, Sweden, Switzerland = commutatus Dieckmann, 1982 gadorensis Stüben, 2001, Spain gracilipes F. Solari, 1938; Algeria (male unknown) guadarramaensis Stüben, 2004; Spain iblanensis Stüben sp. nov.; Morocco kippenbergi Dieckmann, 1982; Italy (l.t.), Switzerland lederi Meyer, 1896; ‘Caucasus’ lemur cisalpinus Stüben, 2003; Italy lemur lemur Germar, 1824 (Cryptorhynchus); Austria, Belgium, France, Germany (l.t.), Italy, Luxembourg, The Netherlands, Switzerland = sulcatus Boheman, 1844 maraoensis Stüben, 2001; Portugal (l.t.), Spain micros Dieckmann, 1982; France, Germany (l.t.), Italy, Switzerland milleri Reitter, 1883; ‘Caucasus’ misellus Boheman, 1844; Denmark, France, Germany, Great Britain (l.t.), Ireland, The Netherlands, Spain, Sweden = variegatus Stephens 1831 [nomen oblitum] = variegatus Stephens 1829 [nomen nudum] monasterialis Stüben, 2004; Spain papei A. & F. Solari, 1905; Bosnia Herzegovina, Croatia, Greece, Hungary, Italy (l.t.), Montenegro, Romania, Serbia = balcanicus A. & F. Solari, 1905 = vicarius Daniel, 1906 parasierrae Stüben, 2002; Morocco parvulus parvulus Boheman, 1837; Austria, Croatia, France (l.t.), Germany, Italy, Slovenia, Switzerland = nudiusculus Foerster, 1849 = turbatus Boheman, 1844 parvulus temperei Péricart, 1987; France (l.t.), Italy, Switzerland petryszaki Dieckmann, 1982; Bulgaria, Poland, Slovakia (l.t.) ptinoides Marsham, 1802 (Curculio); Belgium, Denmark, France, Germany, Great Britain (l.t.), The Netherlands, Portugal, Spain, Sweden = nocturnus Boheman, 1837 reitteri Meyer, 1896, ‘Caucasus’ sardiniaensis Stüben, 2001; Italy (Sardegna) sarothamni Stüben, 2003; Spain setulipennis Desbrochers, 1871; France (Corsica) sierrae H. Brisout, 1865; Spain = bazaensis Stüben, 2001 syn. nov. sintraniensis Stüben, 1999; Portugal



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testensis Stüben, 2003; Morocco tibialis Weise, 1891 (Trachodius); France, Italy (l.t.), Switzerland vorsti Stüben sp. nov.; Spain (Mallorca) Subgenus: Origoacalles Stüben & Astrin, 2010; type species Acalles globulipennis Wollaston, 1854 globulipennis Wollaston, 1854; Portugal (mainland & Madeira, l.t.), Spain (Canary Islands: Gran Canaria, Tenerife) = seminulum Wollaston, 1864: 292 pilula Wollaston, 1864; Spain (Canary Islands: Tenerife, El Hierro, La Palma, l.t.) granulimaculosus Stüben sp. nov.; Spain (Canary Islands: La Gomera) Combinatio nova Coloracalles edoughensis (Desbrochers, 1892) comb. nov. (formerly Acalles); Spain, Algeria (l.t.), Tunisia = hederae Gonzalez, 1965 (Acalles) Pseudodichromacalles xerampelinus (Wollaston, 1864) comb. nov. (formerly Acalles and Canariacalles); Spain (Canary Islands: Tenerife) Incertae sedis Acalles caucasicus Reitter, 1891; Turkey, ‘Caucasus’, Georgia (l.t.). This species may belongs with A. ganglbaueri near A. camelus or into a new genus. Acalles ganglbaueri A. & F. Solari, 1907; Bulgaria (l.t.), Georgia, Greece, Romania, Turkey (see comment regarding A. caucasicus) = ovalipennis Petri, 1912 Acalles pulchellus H. Brisout de Barneville, 1864; France (l.t.), Italy, Spain, Tunesia (probably a species of the genus Kyklioacalles Stüben, 1999) Acalles granulicollis Tournier, 1875 North Africa: Morocco (type missing)

Material and methods The molecular analysis was carried out on 228 Cryptorhynchinae samples: 216 Western Palaearctic, 3 from Mexico and 9 from the United States of America or Canada (currently classified as Acalles, one of them moved to Acallocrates within this work). The 216 Western Palaearctic ones include 177 Acalles s. str. with 37 of 47 known Acalles species / subspecies and 39 Cryptorhynchinae from different genera including the winged Cryptorhynchus lapathi as outgroup species. See Appendix for detailed collecting and vouchering information and GenBank accession numbers. Voucher specimens and extracted genomic DNA are deposited at the Biobank of the Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany (ZFMK). The laboratory routine followed Astrin & Stüben (2008). PCR primers were taken from Astrin & Stüben (2008), CO1 is based on the Folmer et al. (1994) region, 16S is based on the Crandall & Fitzpatrick (1996) region. Both primer sets have been slightly modified for enhanced performance with Cryptorhynchinae weevils. CO1 fw (LCO1490-JJ) 5'-CHACWAAYCATAAAGATATYGG-3' and CO1 rev (HCO2198-JJ) 5'-AWACTTCVGGRTGVCCAAARAATCA-3' 16S fw (16S-1472-JJ) 5'-AGATAGAAACCRACCTGG-3' and 16S rev (16S-ar-JJ) 5'- CRCCTGTTTATTAAAAACAT-3' DNA sequence alignments for concatenated CO1 and 16S datasets were performed with Muscle plugin of Geneious 6.1.6 with default parameters. Primer sequences were trimmed before. Missing data (one gene or nonsequenced nucleotides in the beginning or end of a sequence) were filled up with “n” in the dataset. For all Western Palaearctic species CO1 sequence was available, but 16S sequence data was not available for twelve species: Acalles camelus (I-637-cam), Acalles dubius (D-0094-dub), Acalles kippenbergi (460-PST), Acalles misellus (684-PST), Acalles sardiniaensis (IT1074c), Acalles sardiniaensis (IT1087c), Acalles sardiniaensis (IT1080c), Acalles sierrae (1310-PST), Acalles sierrae (1316-PST), Acalles sierrae (1330-PST), Acalles parvulus temperei (F-804-tem), Calacalles bandamaensis (7-PST). For New World Acalles, 16S data was


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available for all 8 specimens, but CO1 was only available for 3 species: Acalles minutissimus (1763-BAN), Acalles sylvosus (1765-BAN) and Acalles nov. spec. 4 (1767-BAN). The concatenated alignment shows 1229 positions (CO1:658nt, 16S:571nt incl. gaps). The poorly aligned blocks or positions have been determined with GBLOCKS (Castresana 2000, Talavera & Castresana 2007) by using three additional options for less stringent selection: allow smaller final blocks, allow gap positions within the final blocks, allow less strict flanking positions. 1150 nucleotide positions have been used for phylogenetic analysis. The 16S data contained 79 positions, which have been excluded in further calculation. For Bayesian analysis the nucleotide substitution model GTR+I+G (Lanave et al. 1984) has been used for CO1 and 16S partition. We ran MrBayes version 3.2.0 (Ronquist & Huelsenbeck 2003) in two independent replicates, each with 1 cold chain and 3 chains of different temperature (standard setting). For CO1 character sets genetic code for metazoan mitochondrial DNA was used. The third codon position of CO1 and 16S character sets were unlinked in shape, revmat, statefreq and pinvar. The analysis was run for 40 million generations, sampling 40.000 trees. 50.000 generations (which are respectively 50 trees) were discarded for burn-in. Of the retained 39.950 trees a 50%majority-rule consensus tree with posterior probabilities was built (Fig. Acalles tree). Trees were visualized using FigTree v.1.3.1 (Rambaut 2012).

Discussion and results Acalles tree The phylogenetic reconstruction of 37 Western Palaearctic Acalles species and subspecies based on a Bayesian analysis, in general confirm the systematic classification based on morphological characters (Bahr & Stüben 2002). Due to a lack of material, eight species could not be included in the molecular analysis: A. gracilipes, A. granulicollis, A. petryszaki, A. setulipennis. The following four species from the Caucasus have never been collected by the ZFMK or Curculio Institute: A. caucasiscus, A. reitteri, A. milleri and A. lederi. From our point of view, the classification of these four species to Acalles s. str. is questionable. The systematic position of Acalles pulchellus is not clear as well. Only a few specimens have been found so far and because of its specific habitus and distinct internal sac structure we include this species in the genus Kyklioacalles for the time being. Acalles ganglbaueri—one of 'the rare black species'—is mainly distributed in Bulgaria with some localities in Romania. The appearance of A. ganglbaueri is similar to A. caucasicus, but lacks the tuberculate elytral flanks of A. caucasicus. Both might be related to A. camelus, however they may constitute a separate genus, but this cannot be determined at the present time without molecular data. This circumstance certainly applies to A. edoughensis as well (fig. 1a, 1b). This species, which has been collected in Tunisia and Spain around Barcelona, does not belong to Acalles s. str. This has been published in a West-Palaearctic overview tree of Cryptorhynchinae species in Astrin & Stüben et al. (2012) (fig. 3). The nowadays disjunct distribution of Coloracalles humerosus and the distant sister taxon A. edoughensis goes back to the well known land bridge between Sicily and the Iberian Peninsula and North Africa. This is why both species can be found in Algeria on Mt. Edough and in Spain around Barcelona. However, despite the remarkable pdistance of 12.2% of the CO1 gene, which would point to recognition of a separate genus, we think we have found similarities in the elytral marks and endophalli. To avoid another monotypic genus beside Montanacalles nevadaensis, we place Acalles edoughensis in the genus Coloracalles, but the three species Coloracalles edoughensis, C. humerosus and Montanacalles nevadaensis are genetically and morphologically not closely related (see Astrin & Stüben 2008). Pseudodichromacalles xerampelinus also does not belong to Acalles s. str. After one-and-a-half decades of intensive searching for this species, it was finally rediscovered in the winter of 2012 within the laurel forest of Tenerife on the fern Woodwardia radicans (Stüben & Schütte 2013b), 150 years after Wollaston described the species in 1864. At first it seemed impossible to give P. xerampelinus a definite position within the known genera of Cryptorhynchinae from the Canary Islands. After studying the holotype we initially chose the species Canariacalles alluaudi because of the similarity of the internal sac structure of the aedeagus (Stüben 2000c, Stüben & Astrin 2010a: 69). Only the molecular data of CO1 gene made a preliminary assignment possible, namely that it is related to the Pseudodichromacalles species P. fernandezi (Roudier, 1954). Pseudodichromacalles fernandezi is MOLECULAR SYSTEMATICS OF ACALLES


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widespread on El Hierro, La Gomera and Tenerife. In comparison with P. xerampelinus it shows a much wider range of host plants and prefers the sparse and rocky slopes of the laurel forest (Stüben & Astrin 2010a, fig. 9). Taking the elongate habitus and elevated elytral intervals 1 and 3 into account, P. xerampelinus seems to have some morphological similarities to P. fernandezi (fig. 2a). Although the molecular distances are large and sufficient to justifiy a separate genus (10.3%), it seems more appropriate for us to waive the description of a new monotypic genus and keep P. xerampelinus within Pseudodichromacalles at this point (Stüben & Schütte 2013b).

FIGURE 1. Coloracalles edoughensis, habitus and aedeagus.

FIGURE 2. Pseudodichromacalles xerampelinus, habitus and aedeagus.

Acalles gracilipes F. Solari, 1938 from Algeria (only a female holotype available so far) and A. petryszaki Dieckmann, 1982 certainly belong—using morphological aspects—to Acalles s. str. So far there has been no opportunity to retrieve tissue with sufficient quality for sequencing (only an old dry specimen of A. petryszaki was available for sequencing, but unsuccessful). The placement of A. setulipennis Desbrochers, 1871 within Acalles s.str is highly questionable, since its aedeagus has an exceptional ring-like internal sac structure (digit 30: 'Key to the species of Acalles'). This character is more similar to the genus Kyklioacalles Stüben, 1999. The descriptions of Acalles vorsti sp. nov. of the Balearic island Mallorca and Acalles iblanensis sp. nov. of the Middle Atlas mountain range in Morocco are based on "morphological findings". However, we became aware of a new species from the Canary Island La Gomera—Acalles granulimaculosus sp. nov.—after the molecular analysis


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of species belonging to the subgenus Origoacalles. Herewith, Acalles granulimaculosus sp. nov. (type locality: La Gomera) is separated from Acalles pilula Wollaston, 1864 (type locality: La Palma). Although the p-distances of 11.3% to 12.8% (CO1, uncorrected) between these two species leave no room for doubt, there are only a few morphological characters to justify their separation (see Chapter 6, "Taxonomy"). So far this new endemic species is only known from the Canary Island La Gomera, while its sister species A. pilula has spread all over the Canaries. Regarding the species complex A. parvulus/temperei, we refer to our last biogeographical, morphological and molecular studies from 2006: there exists either a hybrid zone, or more likely, that "Acalles parvulus and Acalles temperei constitute a single, geographically structured species" (Stüben & Astrin 2006). The further three species complexes are discussed in detail.

Acalles echinatus species complex Large p-distances at high morphological variability The Central and Eastern European species Acalles echinatus (Germar, 1824) is the sister taxon to the Atlantic species Acalles misellus Boheman, 1844. Both belong in a higher-level clade with Acalles fallax Boheman, 1844 and the type species of the genus Acalles, Acalles camelus (Fabricius, 1792). Acalles camelus falls—which was morphologically expected—in the center of the Acalles species. Also assigned to the Acalles echinatus species complex—but more distant—are Acalles gadorensis Stüben, 2001 (Southern Spain) and Acalles biokovoensis Stüben, 2008 (Croatia). Contrary to the assumption of Lachowska et al. 2009 (based on karyotype studies), Acalles fallax Boheman, 1844 does not belong to the Acalles echinatus complex. By requiring that the genus Acalles follows phylogenetic systematics, the classification of L. Dieckmann (1982) is correct regarding the Acalles echinatus group (with variable species such as A. micros, A. fallax = A. commutatus and maybe A. petryszaki), but it is difficult to maintain his disposition of an Acalles parvulus group (with A. dubius and A. misellus). Also, when comparing the types of Acalles echinatus (Germar, 1824) and Acalles echinatus var. squamosus A. & F. Solari, 1907 syn., in 2003 the second author came to a different evaluation than L. Dieckmann. Dieckmann claimed the aedeagus ("identische Penisform", Dieckmann 1982: 199) of the two species have an identical shape but in fact, the shapes of the aedeagi of specimens from the type locality (Carinthia) and west of Slovenia differ significantly from the ones from France (for example Haute Savoie, Sixt, Nambride, 850 m; Stüben et al. 2003). The aedeagus of those from France is, as aptly depicted in the drawing by A. & F. Solari (1907: 535), much longer and lancet-shaped, while the specimens from the type locality and areas located nearby are shorter, wider and more rounded on the sides. The French specimens might be a separate subspecies of Acalles echinatus, namely Acalles echinatus squamosus A. & F. Solari, 1907. Acalles echinatus seems to be a polymorphic species. We also disagree with Dieckmann’s observation that the shape of the penis, even with material from different geographical areas, is fairly constant („die Form des Penis ist auch bei Material unterschiedlicher geographischer Herkunft ziemlich konstant“, Dieckmann 1982: 205). This observation is based on an insufficient number of specimens, but we agree with his observation of the high variability in appearance of the exoskeleton. The scales on the pronotum can vary from being rounded and smooth to being pointed and more narrow. The lateral punctures of the elytra can be larger and deeper, but also smaller and less deep. The erect bristles on the elytral intervals can be wider and shorter or significantly longer and narrower. This extraordinary variability in the outer appearance of Acalles echinatus is supported by 13 DNA sequences, derived from specimens collected between northern Italy and Moscow. Even though there is a high intraspecific variance between these species, there is no clear speciesspecific clade. The CO1 p-distance range between different A. echinatus populations is 1.4% to 9.1%. The wide distribution area ranges from southern Sweden to Albania and from the French Jura region to the Caucasus. Since this area is far more extensive than the current coverage by molecular data, larger p-distance values have to be assumed by increasing the sampling rate. So far we doubt it might be possible to find sufficient molecular or morphological characters for a species separation by taxonomy or DNA barcoding with CO1. This also applies to other genes like mitochondrial 16S or nuclear gene 28S (Astrin et al. 2012). Extensive cross-breeding might be a promising approach (Stüben 2005). Reliable distance values for species delineation (for example based on CO1 barcoding sequences) are not so easy to set up for the few Cryptorhynchinae with a very large distribution area. Obviosly it's different when dealing



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with flying insects (easier to reach the common gene pool for them), or with widely distributed flightless Cryptorhynchinae. In this case, one must expect not only 'morphological clines' along ecological parameters (e.g. habitat), but also with very large p-distance values of the CO1 gene. Like in the previously handled species complex A. parvulus / A. temperei, here it is very likely that the genetic exchange between subpopulations has stopped and genetic differences (mutations) are not being homogenized anymore (Stüben & Astrin 2006). For a comprehensible assessment of species and species complexes, two steps might point us in the right direction: first, specific CO1 distance values for reliable species delineations for each Curculionidae subfamily; and second, an algorithm that broadly estimates 'relief and distribution' of the specimen of interest: a 'molecular map'. This would be a worthwhile research area which would take into account the possibility that flightless insects, separated by mountain ranges or huge rivers (making use of geological maps), might have not evolved in the same manner that flying ones did during the evolutionary young history (in Western Palearctic, 27 million years ago, Stüben & Astrin 2010: fig. 1B).

Acalles maraoensis species complex Small p-distances but high morphological distinctness At a first glance the four species of the Acalles maraoensis clade offer a uniform appearance like all other species that belong to the Acalles sierrae group of the Spanish and Northern African high mountains (fig. 7; couplets 32–41, ‘Key to the species of Acalles’). These montane open-landscape species have long, oval, egg-shaped, rarely obovate elytra. The intervals of the elytra are much broader than the striae. The integument is consistently dark brown in most species, occasionally with a faded, not rich in contrast, beige spot of scales in front of the base and / or with an indicated fascia behind the middle of the elytra. The elytral bristles are placed distantly in a single row, often at a distance 2 to 3 times bristle length. There are no tufts of bristles like many central European forest species. In this respect, the bristle structure is similar to that of Echinodera. The pronotum is nearly always dark brown and laterally bulbous and rounded, and it is therefore clearly distinct from the broader elytra, which are 1 to 2 intervals wider than the pronotum. The eyes are circular. The rostrum in both sexes is strikingly narrow, and it is approximately as long as the pronotum in the females. Differences in the previously described exoskeletal traits are not extensive or overlap with intraspecific variability within these species. A reliable feature for determination of species of the Acalles maraoensis clade (A. sarothamni, A. monasterialis, A. cytisi and A. maraoensis) is the shape of the median lobe (apex of aedeagus) and the internal sac structure of the aedeagus (couplets 32-43: ‘Key to Acalles’), and for A. cytisi, by using the apically split mid-tibia spine. At least for male specimens, morphological assignment to one of the known species is thus quite possible based on the aforementioned characteristics. According to the molecular results of the CO1 gene analysis, species differentiation is not possible. The phylogenetic reconstruction (Tree 1, lower part) depicts a paraphyletic conflict with the A. maraoensis group. Within the last few years these results have been checked repeatedly—genetically and morphologically. For Cryptorhynchinae systematics, the mitochondrial CO1 gene provided reliable results in virtually 98.5% of the cases, but it is not sufficient for species differentiation within the A. maraoensis group. This is a rare case, but also not the only one in which CO1 fails (Bourke et al. 2013). A similar inconsistency between the morphological and molecular method, can be observed between A. testensis and A. asniensis. These two species from the Morcoccan high mountains do not show extensive exoskeletal differences, but have a totally different internal sac structure of the aedeagus (digit 35). The p-distance between these two species is just 3.2% for CO1 gene and 0.9% for the 16S gene. These values are necessary to evaluate the situation described below. Also, the newly described species Acalles iblanensis can be found within the Moroccan high mountains and has an apically split mid-tibia spine. Acalles iblanensis is the sister species to A. testensis and A. asniensis. Not only does the aedeagus show significant differences (couplets 34, 35), but also the p-distances of A. iblanensis relative to its sister species are at least 11.2% for the CO1 gene and 2.2% for the 16S gene, much larger than the examples cited above. Based on these facts one may assume we are on the safe side for a new species differentiation and description.


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Acalles sierrae species complex Huge p-distances but little morphological distinctness The type locality of Acalles bazaensis Stüben, 2001 is located in the Sierra de Baza (Santa Barbara) in Spain. During the snowmelt in mid-May 2013 we were able to sieve this species under Astragalus. Like the other species of the Acalles sierrae group, this species also prefers Fabaceae host plants. To our surprise, the CO1 sequence of Acalles bazaensis was virtually identical to a specimen found in the eastern Sierra Nevada in 2005 under Erinacea anthyllis. Because of a slightly wider aedeagus, distinct elytra bristles and shorter elytra (digit 41), the eastern one was determined to be Acalles sierrae Brisout, 1865. The formerly depicted differential characters are therefore certainly not species-specific. It has been found that A. bazaensis is a junior synonym of Acalles sierrae, which has been described based on specimens from the Sierra Nevada range. In fact, it is likely the Acalles sierrae species of the Sierra Nevada (not only there) consists of a complex of morphological closely related (sub-) species, that are virtually indistinguishable from each other. For example, a western A. sierrae variant of the Sierra Nevada shows a long rostrum (Stüben 2009: 105) which also applies to specimens that were collected by us just 6 km west of the short rostrum variant on the Puerto de la Ragua mountain. The rostrum length of the Puerto de la Ragua specimen is similar to the ones from Sierra Baza, but the p-distance of the CO1 gene shows at least 6.1% (1316-PST and 1318-PST from Sierra Baza vs. E-0261-sie from Puerto de la Ragua). Specimens of the Acalles sierrae complex of the more distant mountain ranges, namely Sierra La Yedra, Sierra Almadén, Sierra Magina and Sierra de la Pandera, show similar large CO1 p-distances (Tree 1, lower part). These might have slightly wider elytra, but do not provide sufficient morphological characters for species delineation. The differences in the structures of the inner sac of the aedeagus are usually easily comprehensible and a speciesspecific feature. However, in this case they are marginal. They consist of two stripes, more or less parallel in position. We need to accept the fact that the Acalles sierrae species complex (see key below) cannot be resolved based on morphological characters or standard barcoding genes (CO1 and 16S). Further population genetic studies are in preparation. All of these very similar looking variants live in high altitudes—usually above 1300 m—and in isolated mountain massifs of the Betic Cordilleras. They are not found in the plains between the mountain ranges (at least anymore). There could be two possible reasons for this: 1) Under similar environmental conditions, similar structures could have been formed (convergence) or (and more likely), 2) there was not enough time for morphological speciation. In contrast, island species of the Canary Islands and the Madeiran Archipelago show high morphological diversity and very low molecular differences.

Are keys for the identification of insect species still up to date? DNA barcoding put into perspective We are very often asked whether text-based dichotomous determination keys are still relevant. Today, identification courses in the biological sciences at European universities are few or non-existent. Given this distressing state of affairs, it is clear that the days of biological specialists and morphologists seem to be coming to an end. Who will be able to offer correct identifications in the future? The previous process of determination by comparison of morphological features will likely be substituted in a few years by cost-efficient next generation sequencing methods within the framework of DNA barcoding activities. That is why for several years now, most taxonomists and systematists have focussed their work on integrative taxonomy (Dayrat 2005, Will et al. 2005, Padial et al. 2009, Tan et al. 2010, Astrin & Stüben & Misof et al. 2012, Stüben & Astrin 2012). Is this change from the “phenotypic” to the “genotypic” mode of determination already decided? We don't think so and believe that it is worthwhile to invest in pictorial keys, and not only because there is a connection to the above-mentioned insight into the morphology of insects (Klausnitzer 2010: 104/5). There are a lot of subject-didactic solutions based on images for all aspects of classical entomo-morphology. The 'more or less' in elaborate descriptions and comparisons of characteristics can be documented more quickly and impressively by differential pictures, which allow a direct comparison with the insect and its structures under the microscope. That these pictures will be replaced by a modern image recognition system, based on largely automated software—comparable to the DNA barcoding for the (re)identification of species—is likely a question of 'algorithm and time'. The same is true concerning our type collections, which are increasingly difficult to



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maintain in financially strapped European museums with few staff. Digital, high resolution 3D-pictures of habitus, aedeagus and the female reproductive system will reduce the necessity for the loan of specimens and therefore protect the types and reduce the risks associated with shipping. It is true that ‘histories or stories’ do not exist for barcodes and the molecular phylogenetical dendograms that are derived from these [Sudhaus 2006]. Pictures, once again, play an important role in documenting morphological and functional characteristics during evolution (e.g., in the search for homologies). The restructuring of an insect’s mouth into a proboscis could only be possible after the antennae were cleaned with the front legs and not just with the mouthparts. Legs with cleaning bristles or gaps of the fore-tibiae were formed repeatedly and developed independently from one another. Neither barcodes nor molecular phylogenetic reconstructions alone allow the correct perspective. Pictures are also a necessary part of understanding the functional morphology. But at the moment let us return our attention to something different: the applicability of both morphological and molecular methods in entomological science. Whether we call the combination of CO1 sequencing analysis (barcoding), morphological brief descriptions and high-resolution images (Riedel et al. 2013) ‘turbo-taxonomy’ or simply ‘Integrative Taxonomy’ is only of secondary importance. Based on past experience of the editorial staff and their authors of the electronic SNUDEBILLER Journal of the CURCULIO Institute (since 1999), historical type material will not be able to be analysed in the foreseeable future with sensitive and reasonably priced DNA methods. But DNA barcoding must ensure a connection to the well-known and familiar binominal names from the literature. The confidence to correctly identify something is good but what assurance is there that sequences deposited in GenBank indeed correspond to the correct binomial name. There must be a tangible bridge between “genotype and phenotype”, if we do not intend to outdate a 200-year-history of science in entomology. This bridge, to be used by morphologists at any time, is a great challenge (compare it with the digital keys in our electronic journal of the SNUDEBILLER 3/2002, 7-14/2006-2014, the „Digital Weevil Determination“ or DWD project). Up to now a key for identification that includes all Palaearctic Acalles species does not exist. It also seems to be impossible to illustrate and visualize the few existing characteristics of the external skeleton. Fortunately, the digital, web-based stacking images of today allow us to do just this and are available in all imaginable resolutions and they are much more readily obtainable and accurate than any subjective illustration, for example an ink drawing (Stüben, A. 2011).

Key to the species of Acalles Schoenherr, 1825* *Nota: Acalles gracilipes F. Solari, 1938 is not included in the key because males of this species from Algeria are as yet unknown. The suture stripe is treated separately and is not belonging to the (as a rule "only" nine) elytral intervals!


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KEY 1–6.



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KEY 7–13.


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KEY 14–18.



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KEY 19–24.


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KEY 25–29.



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KEY 30–34.


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KEY 35–40.



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KEY 41–43.

Taxonomy Family: Curculionidae Latreille, 1802 Subfamily: Cryptorhynchinae Schoenherr, 1825 Genus: Acalles Schoenherr, 1825, type species: Curculio camelus Fabricius, 1792 New species of the subgenus Acalles s. str. Acalles iblanensis Stüben sp. n. (Figs. 3–7) Type material. Holotype. ♂: “Morocco, M. Atlas, N Imouzzer-des-Marmoucha, Jbel Bou Iblane: Tizi-Bou-Zabel, 33°38'44”N 04°09'18”W, 2275 m, 21.May 2011, Bupleurum, Astragalus, leg. Stüben (44)”, coll. CURCULIOInstitut, D-Mönchengladbach. Paratypes. 11♂, 13♀: data as for holotype, coll. Stüben, CURCULIO-Institut, D-Mönchengladbach; 1♀: “Morocco: Middle Atlas, Jbel Bou Iblane, 2279 m, 33°32’736N 04°09’295 W, 11.5.2009, R. Borovec lgt.”, coll. Borovec. DNAtypes. ♂: data as for holotype, coll. ZFMK: Collectors no: 186-PST_44MO2011, ZFMK-DNA100438120, ZFMK-TIS-100440232; GenBank Acc. no CO1: KC783816, 16S: KJ720538.


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Differential diagnosis. This new species from the Central Atlas (Morocco) belongs to the Acalles sierrae group (see above) of the subgenus Acalles s. str. It is—from a morphological and molecular perspective—closely related to Acalles asniensis Stüben, 2003 and should be compared with the holotype of this species (coll. CURCULIO-Institute) from the type locality near Asni (High Atlas): Acalles iblanensis sp. nov. 1. Clubbed bristles on the elytral intervals are at most 2.5x as long as; their distances range from two to three times the length of a bristle (digit 34). 2. Elytra slightly obovate; behind the middle acuminate-ovally rounded towards the apex (Fig. 3). 3. Apex of the aedeagus (ventral view) elongate and smaller, endophallus with two lines (Fig. 5). Acalles asniensis 1*. Clubbed bristles on the elytral intervals at least 3x as long as, placed much closer (digit 35’). 2*. Elytra broader, laterally short-ovally rounded, and broadly rounded towards the apex (couplet 34 vs. 34’, 35’). 3. Apex of the aedeagus (ventral view) broader; endophallus has a differently shaped structure (digit 35’). The aedeagus of the new species is very similar to the aedeagus of Acalles almeriaensis Stüben, 2001, a widespread species from the south-west of Spain. But the new species from Morocco does not seem to be closely related to this species (couplet 34 vs. 24). Acalles iblanensis sp. nov. 1. Species: 2.3–3.2 mm (without rostrum). 2. Slender and clubbed bristles on the elytral intervals 2.5x as long as wide (couplet 34). 3. Pronotum widest in the middle or immediately behind the middle (digit 34). Acalles almeriaensis 1*. Species smaller: 1.9–2.7mm (without rostrum). 2*. Shorter, wider and more parallelsided bristles on the elytral intervals at most 1.5x as long as wide (digit 24). 3*. Pronotum clearly widest directly in front of the base (digit 24). Description. Length. 2.3–3.2 mm (without rostrum). Head & Rostrum. Eyes large, rounded. Rostrum of males red brown; length-width ratio of 4.0 between insertion of antennae; closely covered with white and beige scales in front of base, finely punctate towards apex; rostrum of females clearly longer; length-width ratio of 5.5 between insertion of antennae, more shiny, more finely punctate. Last five antennal segments as long as broad, trapezoidal, clearly separated from club; first two segments elongate. Pronotum. Only slightly broader than long; widest in middle or immediately behind middle; regularly, strongly rounded laterally towards fore-margin and towards base; with slight depression at sides directly behind fore-margin. In lateral view, contour-lines of pronotum and of elytra form a uniform bend, not clearly separated; in front of fore-margin flattened, towards base slightly more curved. In dorsal view without keels or humps on levelled disc of pronotum. Integument of brown and beige, circular non-overlapping scales, cuticle visible between scales. Small pits of the pronotum are fine, closely separated. Pronotum with very long, slender bristles (dorsal view). Elytra. Length-width ratio of 1.20 (holotype); widest at end of first fourth, slightly obovate; behind middle acuminate-ovally, rounded towards apex. In lateral view the contour-line of elytra flat behind base, forming arc towards apex at declivity. Integument not rich in contrast of (dark)brown and beige, circular non-overlapping scales, cuticle visible between scales. Striae much smaller than intervals, with elongate, closely packed shallow punctures. Clubbed bristles on sutural stripe and intervals at most 2 to 2.5 times as long as wide, their separation ranging from two to three times length of a bristle. Legs. Short; margin of front femur reaching fore-margin of eyes, hind femora ending in front of elytral apex. Covered with predominantly beige, not overlapping, elongate scales and very long, erected bristles, male with split apical spine on middle tibia. Venter. Abdomen with 2nd sternite clearly shorter than 1st sternite and slightly longer than sternite 3 and 4 together.



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FIGURES 3–7. Acalles iblanensis sp. n., habitus and aedeagus of holotype, distribution map of Acalles sierrae group (Spain to Morocco).

Female genitalia. Spermatheca, ovipositor, spiculum ventrale, fig. 6 Aedeagus. Apex of the aedeagus elongate, endophallus with two lines, fig. 5 Etymology. The species name refers to the Mountain of Jbel Bou Iblane (Morocco: Central Atlas). Ecology. The specimens of the type series were sieved under Bupleurum and Astragalus at an elevation of 2300 m a.s.l. Distribution. This species is so far only known from the type locality near Imouzzer-des-Marmoucha, the mountain pass “Tizi-Bou-Zabel” of the Jbel Bou Iblane (Morocco): fig. 7.

Acalles vorsti Stüben sp. n. (Figs. 8–12) Type material. Holotype. ♂: “ES [España: Islas Baleares: Palma de Mallorca] Sóller, Son Coll, N39°46'06" E02°40'12", 14.Apr.2002, 300 m, Quercus forest, leg. O.Vorst, [Es212A]“, coll. CURCULIO-Institut, DMönchengladbach. Paratypes. 2♂, 2♀: data as for holotype, coll. Stüben, coll. Vorst, coll. CURCULIO-Institut, DMönchengladbach. DNAtype. ♀: data as for holotype, coll. ZFMK: Collectors no: 1358-PST_22183, ZFMK-DNA-100417602, ZFMK-TIS-22183; GenBank Acc. no CO1: KF680256, 16S: KJ720563


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Differential diagnosis. The new species from the north of the Balearic Island of Mallorca belongs to the subgenus Acalles s. str. It is, from a morphological and molecular perspective, closely related to Acalles breiti A. & F. Solari, 1909 and should be compared with the lectotype of this species (a male, designated by Stüben 1999, coll. Solari: Museo Civico di Storia Naturale di Milano) from the type locality Valldemossa—only 6 km east of the type locality of the new species (Son Coll): Acalles vorsti sp. nov. 1. Elytra short and oval; widest at or directly in front of the middle (fig. 8, 9). 2. Pronotum widest at the end of the first third in front of the elytral base. 3. Apex of the aedeagus (ventral view) spoon-like, laterally subapically more curved (fig. 10). Acalles breiti 1*. Elytra obovate; widest directly behind the base (digit 27). 2. Pronotum widest in the middle or immediately behind the middle; stronger rounded laterally towards the foremargin than to the elytral base. 3*. Apex of the aedeagus (ventral view) tapered; laterally subapically rectilinear (digit 27). The internal sac structures of the aedeagus of the new species is very similar to the aedeagus of the closely related species Acalles breiti (fig 10 vs. digit 27), but based on our knowledge of more than 1000 sequences of flightless Cryptorhynchinae the CO1 p-distance (uncorrected) of 8.4 to 8.6% between A. vorsti and A. breiti lies above the barcoding gap.

FIGURES 8–12. Acalles vorsti sp.n., habitus and aedeagus of holotype, finding spots for A. vorsti and A. breiti (Mallorca Island). MOLECULAR SYSTEMATICS OF ACALLES


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Description. Length. 2.1–2.9 mm (without rostrum). Head & Rostrum. Eyes small, slightly oval. Rostrum of males deeply punctate, red brown; length-width ratio of 3.5 between insertion of antennae; closely covered with beige scales in front of base, finely punctate at middle, wider, deeply punctate on sides; rostrum of females longer; length-width ratio of 3.8 between insertion of antennae and more shiny (digit 29). Last three antennal segments as long as broad, last segment spherical, other segments trapezoidal, clearly separated from very large, wide club; first two segments elongate. Pronotum. Slightly broader than long; widest at end of first third in front of elytral base; regularly rounded laterally towards fore-margin and towards base; with depression at sides directly behind fore-margin. In lateral view contour-line of pronotum and of elytra clearly separated: pronotum in front of fore-margin flattened, more curved towards base. In dorsal view without keels or humps on flat disc of the pronotum. Integument beige, circular non-overlapping scales, cuticle visible between scales. Small pits of pronotum fine and closely separated. Pronotum with some short, bright, slanted bristles. Elytra. Length-width ratio of 1.20 (holotype); oval, broadest at or directly in front of middle. In lateral view contour-line of elytra flat behind base of declivity forming arc towards apex. Integument of elytra not rich in contrast and consists, like scales of pronotum, of brown and beige circular scales. Striae narrower than intervals, with elongate, shallow, closely packed punctures. Clubbed bristles on sutural stripe and intervals at most 1.5 times as long as wide, their separation ranging from two to five times length of a bristle. Legs. Long; margin of front femur reaching fore-margin of eyes, hind femur ending just in front of elytral apex. Covered with predominantly beige, non-overlapping and very long, close-lying bristles. Venter. Abdomen with 2nd sternite clearly shorter than 1st sternite, slightly longer than sternites 3 and 4 together. Covered with some very long, separate standing bristles. Female genital. Ovipositor, spiculum ventrale, fig. 11 Aedeagus. Apex of the aedeagus (ventral view) spoon-like, laterally subapically more curved, fig. 10 Etymology. I dedicate the new species to my colleague Oscar Vorst (The Netherlands: Utrecht). Ecology. The specimens of the type series were sieved under Quercus. Distribution. So far this species is only known from the type locality near Sóller (Son Coll) on the northwest side of the Balearic island of Mallorca: fig. 12.

New species of the subgenus Origoacalles Acalles granulimaculosus Stüben sp. n. (Figs. 13–15) Type material. Holotype. ♂: “E., Canary Is., La Gomera, S Vallehermoso, Los Loros, garden on a river: Castanea sativa (with connection to laurel forest), N28°08'57" W17°16'36", 609 m, 11.Feb.2010, leg. Stüben (66)”, coll. Museo de Ciencias Naturales de Santa Cruz (Tenerife). Paratypes. 15♂, 5♀: all of them from Spain, Canary Islands: La Gomera: Los Loros, Santa Clara, El Cedro (Las Mimbreras), Rosa de Friate, Alto de La Pietra del Gallo, 1998-2011, leg. Stüben, coll. CURCULIO-Institut, D-Mönchengladbach. DNAtype. ♀: “E: La Gomera: S Vallehermoso: La Meseta, ‘La Piedra Encantada’, Ocotea foetens, N28°09'15" W17°17'36", 819 m, 07.Oct.2008, leg. Stüben & Astrin (20)”, coll. ZFMK: Collectors no: E-724-pil, ZFMK-DNAJJ0651, ZFMK-TIS-cE724; GenBank Acc. no CO1: GU988021 Differential diagnosis. The new species from the Canary Island La Gomera belongs to the subgenus Origoacalles. It is, from a morphological and genetic perspective, closely related to Acalles pilula Wollaston, 1864 and should be compared with species from La Palma (type locality) and El Hierro. Acalles pilula also occurs very rarely in the north of Tenerife (digit 43), but is not identical with Acalles pilula var. seminulum Wollaston, 1864 here, which was described also from Tenerife (designation of lectotype see Stüben 2000c, page 84-85). This name is a junior synonym of Acalles globulipennis Wollaston, 1854, a species from Tenerife, Gran Canaria and Madeira (digit 42), which prefers the transition zone between laurel forest and thermophilic brushwood, and can also be found in the low-lying areas near the north coast of Portugal (own records, Stüben, Behne & Bahr 2003).


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Acalles granulimaculosus sp. nov. 1. Pronotum with circular, considerably large scales (digit 43’). 2. Aedeagus (ventral view): base broader; the apex, in lateral view, nearly curved at a right angle (fig. 15). Acalles pilula 1*. Pronotum behind the front margin on center line with predominant elongate-oval and smaller scales (digit 43). 2*. Aedeagus (ventral view): narrower, median lobe very long; the apex, in lateral view, less curved (digit 43).

FIGURES 13–15.

Description. Length. 2.0–2.6 mm (without rostrum). The new species corresponds in all other details—excluding the characteristics listed above, with the sister taxon A. pilula from Tenerife and the western islands El Hierro and La Palma (type locality). A detailed redescription of A. pilula with many scientific illustrations and a comparison with A. globulipennis can be found in Stüben (2000: 82-85). See also the couplet 43 vs. 43’ in this study. Etymology. The species name refers to the small body (latin: granulum) and in many cases soiled, but always colored elytra (latin: maculosus) Ecology. The specimens of the new species can be knocked off dead and broken branches of bushes and trees of the laurel forest and the thermophilic brushwood. Distribution. So far this species is only known from the Canarian Island La Gomera.

Excursus: The New World species of "Acalles" Ch. S. Papp mentioned 119 Acalles species in his work "Cryptorhynchinae of the New World" in 1979. At a first glance these species from North, Central and South America look similar to the Acalles species of the Western Palaearctic. Morphologically, the differences are actually difficult to see using exoskeletal characters but are directly visible from examining the very different inner sac structure (endophallus) of the aedeagus. Actually, the western palaearctic species show different inner sac structures—with one exception (see below). Unfortunately, the apparent assumption that the New World "Acalles" are typical representatives of the genus Acalles (Rheinheimer 2007/2008) is a deception. The outwardly similar appearance indicates an adaptation to similar ecological conditions and leads to convergent development, especially in the leaf litter and wood litter of the American forests. Of the 12 well-known species of the North American "Acalles", 6 named species and 2 undetermined ones have been kindly provided by R. S. Anderson of the Canadian Museum of Nature, Research and Collections Division in Ottawa, Canada. These are: Acalles costifer Le Conte, 1884; A. indigens Fall, 1907; A. minutissimus (Illiger, 1807); A. sablensis Blatchley, 1920; A. sylvosus Blatchley, 1916; A. crassulus Le Conte, 1876; Acalles sp. 4 and Acalles sp. 5. These specimens MOLECULAR SYSTEMATICS OF ACALLES


27

have been handled in accordance with the materials and methods specified above: the DNA barcoding area of CO1 and a part the 16S gene have been sequenced and added to our Acalles dataset for Bayesian analysis. These first molecular results (Tree 1, upper part) for New World Acalles reveal that none of the specimens from the United States are closely related to the type species of the genus Acalles, namely Acalles camelus (Tree 1, lower part). They are not grouped with the flightless sister genera like Echinodera, Dichromacalles and Kyklioacalles or the numerous genera of the Macaronesian Islands. Therefore we recommend that a new genus should be described for these so-called "Acalles" species from North America. We make only one exception: Acalles costifer Le Conte, 1884, which we transfer to the genus Acallocrates Reitter, 1913 (cf. Stüben et al. 2013: Fig 1.). This genus is basally placed in the phylogenetic tree.

FIGURES 16–23. New World Species of "Acalles", habitus and aedeagus of Acallocrates costifer, Acalles crassulus, Acalles sylvosus and Acalles indigens.

Acalles costifer was initially suspected to be an Acallocrates (compare fig. 16, fig. 20). This suspicion was based upon two morphological features: the deep, double curved base of the pronotum and the basal angles protruding from the sides of the elytra (extended shoulder, fig. 16). These features are mainly due to the inner sac


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structure of the aedeagus which is characteristic of Acallocrates (compare with fig. 24, 25 and Bahr 2003). The above mentioned suspicion based on morphological characters has now been confirmed by molecular analysis. However, these first impressions and results that we have stated here we hope to expand upon in the upcoming years in the M.W.I. Project (Molecular Weevil Identification). For this task, we ask colleagues to send us determinated specimens in ethanol from all over the world, from as many Cryptorhynchinae genera as possible.

FIGURES 24–25. Acallocrates minutesquamosus from Europe, habitus and aedeagus.

Acknowledgements This article was prepared by the Molecular Weevil Identification project (MWI) of the CURCULIO Institute (CURCI) & Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Molecular Taxonomy Section. We wish to thank the Spanish and Morroccan administration offices for scientific collection permits. We also extend our gratitude to Dr. Robert Anderson of the Canadian Museum of Nature in Ottawa, Canada, for the set of New World weevils. Lastly, we wish to thank Kevin Smith of Carleton University in Ottawa for his meticulous editing and proof-reading of our manuscript.

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APPENDIX TABLE 1. Collecting data, voucher numbers and GenBank accession numbers for the material analysed in this study for phylogenetic tree of (subfamily Cryptorhynchinae). Collecting dates are given in the form dd.mm.yyyy. Taxon

Collecting Data

ZFMK no. / Collectors no. DNA no. / tissue no. CO1: Genbank acc. no. 16S: genbank acc. no.

Acalles alcarazensis Stüben, 2009

SPAIN: Castilla-La Mancha, Sierra de Alcaraz, 12 km SE Alcaraz, near Mt. Almenara, N38°34'17", W02°25'53", Amelanchier ovalis, Acer, 1650m, 22.04.2003, leg. P.Stüben, det. P.Stüben, col. ZFMK

9129/E-457-alc DNA-JJ0467/TIS-cE457 CO1: GU987921 16S: GU988349







Acalles almeriaensis Stüben, 2001

SPAIN: Almería, 11 km NW Laujar de Andarax, Sierra Nevada, Bayárcal, N37°02'27", W03°00'12", Quercus ilex, Cytisus scoparius, Euphorbia, 1291m, 07.04.2007, leg. J.Astrin and P.Stüben, det. P.Stüben, col. ZFMK

8836/E-0186-alm DNA-JJ0174/TIS-cE0186 CO1: EU286497 16S: EU286333


SPAIN: Granada, 5 km N Laujar de Andarax, Sierra Nevada, N37°02'16", W02°54'51", Erinacea anthyllis, 1597m, 07.04.2007, leg. J.Astrin and P.Stüben, det. P.Stüben, col. ZFMK

8921/E-0192-alme DNA-JJ0259/TIS-cE0192 CO1: GU987825 16S: GU988242


SPAIN: Teruel, S. Javalambre, Fuente la Risca near Arcos de las Salinas, N39°59'56", W01°01'21", Amelanchier ovalis, Acer monspessulanum, Erinacea anthyllis, Ulex, 1121m, 17.06.2008, leg. J.Astrin, det. P.Stüben, col. ZFMK

9202/E-601-alm DNA-JJ0540/TIS-cE601 CO1: GU987949 16S: GU988389


SPAIN: Teruel, near Mora de Rubielos, N40°19'08", W00°43'19", 9204/E-603-alm Erinacea anthyllis, 1369m, 18.06.2008, leg. J.Astrin, det. P.Stüben, DNA-JJ0542/TIS-cE603 col. ZFMK CO1: GU987951 16S: GU988391


SPAIN: Castellón, Morella, Barranco de la Bota, N40°33'12", W00°00'27", Quercus ilex, Hedera helix, 814m, 21.06.2008, leg. J.Astrin, det. P.Stüben, col. ZFMK



SPAIN: Málaga, NE Málaga, Sierra de Tejeda, Alcaucin, N36°56'13", W04°04'59", Quercus ilex, 1020m, 07.01.2009, leg. Stüben, det. P.Stüben, col. ZFMK



SPAIN: Málaga, NE Málaga, Sierra de Tejeda, Alcaucin, N36°54'54", W04°05'27", Viburnum tinus, 751m, 07.01.2009, leg. Stüben, det. P.Stüben, col. ZFMK



SPAIN: Malaga, S of Fornes: Sierra de Almijara (N. P.), N36°53'21", W03°51'14", Quercus, Viburnum, ähnlich Kreuzdorn, 1002m, 08.05.2013, leg. P.Stüben & A.Schütte, det. P.Stüben, col. ZFMK

10180/1280-PST DNA-100426197/TIS-4096 CO1: KF680246 16S: KJ720556

Acalles asniensis Stüben, 2003

MOROCCO: High Atlas, E Asni, Oukaimeden, N31°12'14", W07°52'21", Erinacea, 2646m, 01.05.2009, leg. P.Stüben, det. P.Stüben, col. ZFMK

9501/E-882-asn DNA-JJ0839/TIS-cE882 CO1: GU988129 16S: GU988569 ......continued on the next page



33

APPENDIX TABLE 1. (Continued) Taxon

Collecting Data


Acalles asniensis Stüben, 2003

9503/E-884-asn MOROCCO: High Atlas, SE Asni, Tacheddirt (near Imlil), N31°09'07", W07°50'18", broom, Daphne, 2420m, 02.05.2009, leg. DNA-JJ0841/TIS-cE884 CO1: GU988130 P.Stüben, det. P.Stüben, col. ZFMK 16S: GU988570

Acalles biokovoensis Stüben, 2008

CROATIA: Dalmatien, 3 km NE Makarska, Biokovo Mts., "Vosac", 8891/HR-0330-bio N43°18'53", E17°03'09", Fagus, 1367m, 11.07.2007, leg. P.Stüben, DNA-JJ0229/TIS-cHR0330 det. P.Stüben, col. ZFMK CO1: GU987819 16S: GU988236

Acalles biokovoensis Stüben, 2008

CROATIA: Dalmatien, 4 km E Makarska, Biokovo Mts., "Mt. Sinjal", N43°18'39", E17°03'49", Fagus, 1320m, 16.07.2007, leg. P.Stüben, det. P.Stüben, col. ZFMK

8892/HR-0341-bio DNA-JJ0230/TIS-cHR0341 CO1: GU987820 16S: GU988237

Acalles breiti A. & F. Solari, 1909

SPAIN: Mallorca, Serra de Tramuntana, 2,5 km NO Puig Major, N39°48'59", E02°49'26", Quercus ilex, 29.12.2007, leg. J.Astrin, det. J.Astrin, col. ZFMK

9104/E-542-bre DNA-JJ0442/TIS-cE542 CO1: GU987911 16S: GU988339

Acalles breiti A. & F. Solari, 1909

SPAIN: Mallorca, Serra de Tramuntana, Sa Calobra, N39°51'04", E02°48'23", Quercus ilex, 27.12.2007, leg. J.Astrin, det. J.Astrin, col. ZFMK

9105/E-543-bre DNA-JJ0443/TIS-cE543 CO1: GU987912 16S: GU988340

Acalles camelus (Fabricius, 1792)

FRANCE: Isère, 2 km SE Lans en Vercors, Montagne de Lans, N45°06'45", E05°36'21", Abies, Fagus, Fraxinus, 1352m, 12.08.2005, leg. P.Stüben, det. P.Stüben, col. ZFMK

8679/I-0026-cam DNA-JJ0017/TIS-cI0026 CO1: EU286447 16S: EU286282


FRANCE: Isère, 14 km N Grenoble, Massif de la Chartreus, NW Col de Porte, N45°18'40", E05°45'17", Abies, Fagus, Fraxinus, 1649m, 13.08.2005, leg. P.Stüben, det. P.Stüben, col. ZFMK

8707/I-0034-cam DNA-JJ0045/TIS-cI0034 CO1: GU987776 16S: GU988186


FRANCE: Isère, 9 km N Grenoble, Mont St. Martin, N45°16'15", E05°40'44", Fagus, Fraxinus, Quercus, 813m, 12.08.2005, leg. P.Stüben, det. P.Stüben, col. ZFMK

8824/I-0016-cam DNA-JJ0162/TIS-cI0016 CO1: GU987808 16S: GU988224


SLOVENIA: Apasko Polje, Podgorje, Podgorje env., 30 km NE of 8845/SLO-0227-cam Maribor; Quercus, Carpinus, N46°42', E15°55', Quercus, Carpinus, DNA-JJ0183/TIS-cSLO0227 05.08.2006, leg. R.Stejskal, det. R.Stejskal, col. ZFMK CO1: GU987813 16S: GU988229


AUSTRIA: Niederösterreich, Merkersdorf, National Park Thayatal; 8846/A-0228-cam Carpinetum, N48°32', E16°17', 18.04.2006, leg. R.Stejskal, det. DNA-JJ0184/TIS-cA0228 R.Stejskal, col. ZFMK CO1: GU987814 16S: GU988230


CROATIA: Dalmatien, 8 km E Karlobag, Velebit Mts., Stupacinovo, N44°32'41", E15°09'58", Fagus, 1049m, 14.07.2007, leg. P.Stüben, det. P.Stüben, col. ZFMK

8856/HR-0336-cam DNA-JJ0194/TIS-cHR0336 CO1: EU286511 16S: EU286347


SLOVENIA: Drava Valley, Cresnjevec, Cresnjevec env., 22 km W of Maribo, N46°32', E15°22', Fagus, Acer, Abies, 04.08.2006, leg. R.Stejskal, det. R.Stejskal, col. ZFMK

8946/SLO-0229-cam DNA-JJ0284/TIS-cSLO0229 CO1: GU987842 16S: GU988259 ......continued on the next page


SCHÜTTE & STÜBEN


Collecting Data



ITALY: Campania, Cilento, 6 km SE Vallo d. Lucania, M. Sacro o 9229/I-628-cam Gelbison, N40°12'41", E15°19'42", Fagus, 1544m, 30.06.2008, leg. DNA-JJ0567/TIS-cI628 CO1: GU987971 P.Stüben, det. P.Stüben, col. ZFMK 16S: GU988411


ITALY: Basilicata, Monte Pollino, 9 km SE Rotonda, Rif. de Gasperi, N39°54'37", E16°07'15", Fagus, 1486m, 05.07.2008, leg. P.Stüben, det. P.Stüben, col. ZFMK


ITALY: Abruzzo, P.N. Majella, 11 km N Roccaraso, Bosco di S. 9257/I-656-cam Antonio, N41°56'27", E14°01'41", Pyrus/Acer, 1321m, 14.07.2008, DNA-JJ0595/TIS-cI656 leg. P.Stüben, det. P.Stüben, col. ZFMK CO1: GU987989 16S: GU988427


SWEDEN: Simrishamn, Stenshuvud N.P., N55°39'15", E14°15'50", 9333/S-662-cam 80m, 12.08.2008, leg. J.Astrin, det. P.Stüben, col. ZFMK DNA-JJ0671/TIS-cS662 CO1: GU988028 16S: GU988465


GERMANY: Rügen, Sassnitz, near Rusewase, N54°32'29", E13°37'46", Fagus, 144m, 15.08.2008, leg. J.Astrin, det. P.Stüben, col. ZFMK

9337/D-666-cam DNA-JJ0675/TIS-cD666 CO1: GU988032 16S: GU988469


CZECH REPUBLIC: SE Moravia (UH), Lopenik, Bilé Karpaty Mts., N48°56', E17°46', from trees of Fagus, 15.05.2009, leg. Krsl, det. Krsl, col. ZFMK

9637/CZ1054 DNA-JJ1054/TIS-cCZ1054 CO1: KF680229 16S: KJ720541

Acalles cazorlaensis Stüben, 2004

SPAIN: Andalucía, Sierra de Segura (Sierra de Pozo), 12 km SE Cazorla, N37°52'53", W02°52'39", Quercus ilex, 1300m, 17.04.2003, leg. P.Stüben, det. P.Stüben, col. ZFMK

9021/E-464-caz DNA-JJ0359/TIS-cE464 CO1: GU987884 16S: GU988305


9197/E-596-caz SPAIN: Jaén, Cazorla, El Chorro, N37°50'25", W02°59'24", Erinacea anthyllis, 1559m, 12.06.2008, leg. J.Astrin, det. P.Stüben, DNA-JJ0535/TIS-cE596 CO1: GU987946 col. ZFMK 16S: GU988386


SPAIN: Jaen, Sierra de Cazorla, S of Cazorla: near El Chorro, N37°50'20", W02°59'26", Erinacea anthyllis, (Euphorbia nicaeensis), 1581m, 16.05.2013, leg. P.Stüben & A.Schütte, det. P.Stüben, col. ZFMK


SPAIN: Jaén, Cazorla, near El Chorro, N37°50'52", W02°57'34", 9200/E-599-caz Erinacea anthyllis, 1610m, 12.06.2008, leg. J.Astrin, det. P.Stüben, DNA-JJ0538/TIS-cE599 col. ZFMK CO1: GU987948 16S: GU988388

Acallocrates costifer comb. nov. (LeConte, 1884)

USA: Texas, Colorado County, Columbus, N29°41', W96°30', 21.02.1989, leg. R.Anderson, det. R.Anderson, col. ZFMK

11972/1761-BAN DNA-0155668051/TIS-24972 CO1: n.a. 16S: KJ767025

Acalles crassulus LeConte, 1876

USA: Florida, Dade County, Miami, Charles Deering Estate Park, N25°36', W80°18', hardwood hammock litt., 16.10.1990, leg. R.Anderson, det. R.Anderson, col. ZFMK


9238/I-637-cam DNA-JJ0576/TIS-cI637 CO1: GU987978 16S: n.a.


......continued on the next page



35


Collecting Data


Acalles cytisi Stüben, 2004

SPAIN: Cáceres, 28 km N Plasencia, Montes de Tras la Sierra, Puerto de Honduras, N40°13'58", W05°53'02", Cytisus, 1273m, 09.04.2004, leg. P.Stüben, det. P.Stüben, col. ZFMK

9019/E-462-cyt DNA-JJ0357/TIS-cE462 CO1: GU987883 16S: GU988304


SPAIN: Ávila, NW Arenas de San Pedro, Gredos NP, "la 9465/E-845-cyt plataforma", N40°16'06", W05°14'22", Cytisus, 1880m, 24.04.2009, DNA-JJ0803/TIS-cE845 leg. J.Astrin, det. P.Stüben, col. ZFMK CO1: GU988104 16S: GU988543


SPAIN: Ávila, E Béjar, S La Hoya, La Covatilla, N40°21'26", W05°41'06", Cytisus, 1950m, 24.04.2009, leg. J.Astrin, det. P.Stüben, col. ZFMK

9467/E-847-cyt DNA-JJ0805/TIS-cE847 CO1: GU988105 16S: GU988544

Acalles dieckmanni Pericart, 1989

FRANCE: Hte Corse, Vallée Restonica, N42°14'10", E09°02'10", 25.05.2011, leg. Lempereur, det. P.Stüben, col. ZFMK


Acalles dubius A. & F. Solari, 1907

GERMANY: RLP, Bausenberg, Niederzissen, N50°27'52", E07°13'29", Quercus, Carpinus, Fagus, 291m, 17.09.2005, leg. P.Stüben, det. P.Stüben, col. ZFMK

8694/M-0066-dub DNA-JJ0032/TIS-cM0066 CO1: GU987767 16S: GU988177


GERMANY: Baden-W., Kaiserstuhl, Auf dem Eck, N48°06'32", E07°40'04", Fagus, Carpinus, Quercus, 432m, 27.05.2006, leg. P.Stüben, det. P.Stüben, col. ZFMK

8822/D-0094-dub DNA-JJ0160/TIS-cD0094 CO1: GU987806 16S: n.a.


GERMANY: RLP, Kaltenborn, Hohe Acht, N50°23'08", E07°01'25", Quercus, Fagus, 605m, 03.11.2007, leg. P.Stüben, det. P.Stüben, col. ZFMK

9009/D-0423-dub DNA-JJ0347/TIS-cD0423 CO1: GU987881 16S: GU988298


GERMANY: Rhineland-Palatinate, E Treis-Karden, Pommern, Pommernbachtal, forest path, N50°10'52", E07°16'42", Fagus, 116m, 09.06.2012, leg. P.Stüben, det. P.Stüben, col. ZFMK


Acalles echinatus (Germar, 1824)

AUSTRIA: Niederösterreich, Merkersdorf, National Park Thayatal, 8737/A-0093-ech N48°32', E16°17', Carpinetum, 18.04.2006, leg. R.Stejskal, det. DNA-JJ0075/TIS-cA0093 R.Stejskal, col. ZFMK CO1: GU987786 16S: GU988200


CROATIA: Dalmatien, 6 km E Karlobag, Velebit Mts., Ostarijska vrata, N44°31'45", E15°08'34", Fagus, 927m, 14.07.2007, leg. P.Stüben, det. P.Stüben, col. ZFMK

8855/HR-0334-ech DNA-JJ0193/TIS-cHR0334 CO1: EU286510 16S: EU286346


GERMANY: Bayern, N Karlstadt, Gambach, NSG GrainbergKalbenstein, N49°59'28", E09°45'45", Fagus, 260m, 19.05.2006, leg. P.Stüben, det. P.Stüben, col. ZFMK

8864/D-0234-ech DNA-JJ0202/TIS-cD0234 CO1: GU987816 16S: GU988232


CZECH REPUBLIC: Moravia mer., Pavlov env, Pálava Protected Landscape Area, N48°52', E16°40', Tilia, Carpinus, Quercus, 28.05.2006, leg. R.Stejskal, det. R.Stejskal, col. ZFMK

8948/CZ-0231-ech DNA-JJ0286/TIS-cCZ0231 CO1: GU987843 16S: GU988260 ......continued on the next page


SCHÜTTE & STÜBEN


Collecting Data



CROATIA: Dalmatien, 8 km E Karlobag, Velebit Mts., Stupacinovo, N44°32'41", E15°09'58", Fagus, 1049m, 14.07.2007, leg. P.Stüben, det. P.Stüben, col. ZFMK

8992/HR-0338-ech DNA-JJ0330/TIS-cHR0338 CO1: GU987870 16S: GU988287


BULGARIA: W Gabrovo, Aprilci, N42°50'26", E24°55'10", 515m, 8993/BUL-0344-ech 08.06.2007, leg. Bialooki, det. Bialooki, col. ZFMK DNA-JJ0331/TIS-cBUL0344 CO1: GU987871 16S: GU988288


BULGARIA: W Gabrovo, Stokite, N42°51'47", E25°04'19", 372m, 8994/BUL-0345-ech 11.06.2007, leg. Bialooki, det. Bialooki, col. ZFMK DNA-JJ0332/TIS-cBUL0345 CO1: GU987872 16S: GU988289


RUSSIA: Ivanteevka, near Moscow, N55°59'14", E37°58'37", Quercus, Fagus, Corylus, 38m, 18.08.2007, leg. J.Astrin, det. J. Astrin, col. ZFMK

9007/RU-0410-ech DNA-JJ0345/TIS-cRU0410 CO1: GU987879 16S: GU988296


RUSSIA: Moskow, near Ivanteevka, N55°59'18", E37°59'00", 170m, 01.09.2008, leg. J.Astrin, det. P.Stüben, col. ZFMK

9339/R-668-ech DNA-JJ0677/TIS-cR668 CO1: GU988033 16S: GU988470


RUSSIA: Moskow, Fryazino, Grebnevo, N55°57'01", E38°05'05", Betula, Tilia, 160m, 02.09.2008, leg. J.Astrin, det. P.Stüben, col. ZFMK

9340/R-669-ech DNA-JJ0678/TIS-cR669 CO1: GU988034 16S: GU988471


ITALY: Monti Berici (Vl), dint. San Gottardo, N45°27', E11°28', 25.02.2009, leg. L.Diotti, det. L.Diotti, col. ZFMK

9632/IT1049 DNA-JJ1049/TIS-cIT1049 CO1: KF680227 16S: KJ720539


SLOVAKIA: Slizké env., N48°29'52", E20°05'28",Quercetum, 284m, 29.09.2012, leg. J.Kratky, det. J.Kratky, col. ZFMK

10121/1221-JKR DNA-100426063/TIS-3557 CO1: KF680241 16S: KJ720551


ITALY: Vicenza, Monti Berici, dintorni Villabalzana, N45°26', 10371/461-PST E11°31', 385m, 26.02.2011, leg. L.Diotti, det. P.Stüben, col. ZFMK DNA-100417598/TIS-20247 CO1: KF680257 16S: KJ767023

Acalles fallax Boheman, 1844

CROATIA: Dalmatien, 22 km SE Gospic, Velebit Mts., Medak, 8853/HR-0332-fal N44°24'32", E15°34'13", Quercus, 617m, 13.07.2007, leg. P.Stüben, DNA-JJ0191/TIS-cHR0332 det. P.Stüben, col. ZFMK CO1: EU286508 16S: EU286344


SLOVENIA: Drava Valley, Cresnjevec env., 22 km W of Maribor, N46°32', E15°22', Fagus, Acer, Abies, 04.08.2006, leg. R.Stejskal, det. R.Stejskal, col. ZFMK

8950/SLO-0235-fal DNA-JJ0288/TIS-cSLO0235 CO1: GU987844 16S: GU988261


CZECH REPUBLIC: W Bohemia (KT), Klenová (6645), N49°20', E13°13', 24.08.2008, leg. Kresl, det. Kresl, col. ZFMK

9425/Cz-797-fal DNA-JJ0763/TIS-cCz797 CO1: GU988078 16S: GU988518 ......continued on the next page



37


Collecting Data



GERMANY: NRW, N Aachen, N Aachen, S. Herzogenrath, N50°51'25", E06°05'44", 262m, 28.03.2009, leg. P.Stüben, det. P.Stüben, col. ZFMK

9451/D-831-fal DNA-JJ0789/TIS-cD831 CO1: GU988093 16S: GU988532


CZECH REPUBLIC: SE Moravia (UH), Lopenik, Bilé Karpaty 9635/CZ1052 Mts., N48°56', E17°46', Fagus, 15.05.2009, leg. Krsl, det. Krsl, col. DNA-JJ1052/TIS-cCZ1052 ZFMK CO1: KF680228 16S: KJ720540

Acalles gadorensis Stüben, 2001

SPAIN: Murcia, 13 km N Totana, Sierra de Espuna, N37°52'07", W01°33'46", Erinacea anthyllis, Quercus ilex under Pinus, 1361m, 29.12.2005, leg. P.Stüben, det. P.Stüben, col. ZFMK

8740/E-0097-gad DNA-JJ0078/TIS-cE0097 CO1: EU286459 16S: EU286294

Acalles globulipennis SPAIN: Canary Islands, Gran Canaria, Moya, Barranco de los Tilos, 204/14-PST_7GC2011 DNA-100438096/TISWollaston, 1854 N28°05'18", W15°35'36", Laurisilva, 446m, 07.12.2010, leg. 2D100447024 P.Stüben, det. P.Stüben, col. ZFMK CO1: JN701882 16S: KJ720537 Acalles globulipennis PORTUGAL: Madeira, 4,5 km S São Vicent Boca da Encumeada, Wollaston, 1854 "Folhadal", N32°45'08", W17°01'40", Laurisilva, Euphorbia mellifera, 1004m, 22.03.2008, leg. J.Astrin and P.Stüben, det. J.Astrin and P.Stüben, col. ZFMK

9092/P-528-glo DNA-JJ0430/TIS-cP528 CO1: FJ716546 16S: FJ716495

Acalles globulipennis PORTUGAL: Madeira, 3 km W Porto Moniz, Santa Madalena, Wollaston, 1854 N32°51'31", W17°12'11", Ficus carica, 391m, 19.03.2008, leg. J.Astrin and P.Stüben, det. J.Astrin and P.Stüben, col. ZFMK

9118/P-521-glo DNA-JJ0456/TIS-cP521 CO1: GU987919 16S: GU988346

Acalles globulipennis SPAIN: Canary Islands, Tenerife, NE La Laguna, Anaga Mts. near 9265/E-676-glo Wollaston, 1854 Moquinal, N28°31'55", W16°17'24", Laurisilva, 840m, 29.09.2008, DNA-JJ0603/TIS-cE676 CO1: FJ716562 leg. J.Astrin and P.Stüben, det. P.Stüben, col. ZFMK 16S: FJ716513 Acalles globulipennis SPAIN: Canary Islands, Tenerife, NE La Laguna, Anaga Mts. near Wollaston, 1854 Chinobre, N28°33'21", W16°10'46", Laurus, Ixanthus viscosus, 808m, 30.09.2008, leg. J.Astrin and P.Stüben, det. P.Stüben, col. ZFMK

9268/E-679-glo DNA-JJ0606/TIS-cE679 CO1: GU987997 16S: GU988435

Acalles globulipennis PORTUGAL: N Caldas da Rainha., SW Nazaré, near São Martinho 9485/E-866-glo Wollaston, 1854 do Porto, N39°30'23", W09°09'16", Pistacia lentiscus, 53m, DNA-JJ0823/TIS-cE866 28.04.2009, leg. J.Astrin, det. P.Stüben, col. ZFMK CO1: GU988117 16S: GU988556 Acalles globulipennis PORTUGAL: Estremadura, NW Sintra: Serra de Sintra, Colares: Pé 9744/843-PST DNA-100413782/TIS-3756 Wollaston, 1854 da Serra, N38°47'13", W09°27'46", Quercus, 323m, 16.05.2012, CO1: KF680240 leg. P.Stüben, det. P.Stüben, col. ZFMK 16S: KJ720550 Acalles granulimaculosus spec.nov. Stüben, 2014

SPAIN: Canary Islands, La Gomera, S Hermigua, El Cedro, Las Mimbreras, N28°07'27", W17°13'26", Laurisilva, 901m, 06.10.2008, leg. J.Astrin and P.Stüben, det. P.Stüben, col. ZFMK

9306/E-717-pil DNA-JJ0644/TIS-cE717 CO1: GU988017 16S: GU988454

Acalles granulimaculosus spec.nov. Stüben, 2014

SPAIN: Canary Islands, La Gomera, S Vallehermoso, La Meseta, "La Piedra Encantada", N28°09'15", W17°17'36", Ocotea foetens, 819m, 07.10.2008, leg. J.Astrin and P.Stüben, det. P.Stüben, col. ZFMK

9313/E-724-pil DNA-JJ0651/TIS-cE724 CO1: GU988021 16S: GU988458 ......continued on the next page


SCHÜTTE & STÜBEN


Collecting Data


Acalles guadarramaensis Stüben, 2004

ANDORRA:, near El Serrat, N42°37'38", E01°33'07", Corylus avellana, Betula, 1800m, 24.06.2008, leg. J.Astrin, det. P.Stüben, col. ZFMK

9218/AND-617-gua DNA-JJ0556/TIS-cAND617 CO1: GU987963 16S: GU988403


SPAIN: Madrid, N Somosierra, Pto. de Somosierra, N41°08'39", W03°34'41", Cytisus, Ilex, Salix, 1380m, 22.04.2009, leg. J.Astrin, det. P.Stüben, col. ZFMK

9456/E-836-gua DNA-JJ0794/TIS-cE836 CO1: GU988096 16S: GU988535


SPAIN: Community of Madrid, NE of Montejo de la Sierra, NE of Hiruela: Hayedo de Montejo, N41°05'23", W03°25'41", Quercus, 1183m, 02.05.2013, leg. P.Stüben & A.Schütte, det. P.Stüben, col. ZFMK



SPAIN: Community of Madrid, N of Lozoya, S of Puerto de Lozoya 10170/1270-PST DNA-100426486/TIS-4086 o de Navafria, N40°58'08", W03°48'00", Cytisus, 1294m, CO1: KF680245 03.05.2013, leg. P.Stüben & A.Schütte, det. P.Stüben, col. ZFMK 16S: KJ720555

Acalles iblanensis spec.nov. Stüben, 2014

MOROCCO: M. Atlas, N Imouzzer-des-Marmoucha, Jbel Bou Iblane: Tizi-Bou-Zabel, N33°38'44", W04°09'18", broom, Bupleurum, Astragalus, 2275m, 21.05.2011, leg. P.Stüben, det. P.Stüben, col. ZFMK

381/186-PST_44MO2011 DNA-100438120/TIS2D100440232 CO1: KC783816 16S: KJ720538

Acalles indigens Fall, 1907

USA: Texas, Culberson County, Guadalupe Mountains National Park: McKittrick Canyon, N31°58', W100°45', 1645m, 10.09.1988, leg. R.Anderson, det. R.Anderson, col. ZFMK


Acalles kippenbergi Dieckmann, 1982

ITALY: Verona, Badia Calavena, frazione RAUSI, N45°34', 664/460-PST E11°09', 500m, 27.02.2007, leg. L.Diotti, det. P.Stüben, col. ZFMK DNA-0100449382/TIS-20246 CO1: KJ867571 16S: n.a.

Acalles lemur cisalpinus Stüben, 2003

ITALY: Campania, Cilento, 6 km SE Vallo d. Lucania, M. Sacro o 9227/I-626-cis Gelbison, N40°12'41", E15°19'42", Fagus, 1544m, 30.06.2008, leg. DNA-JJ0565/TIS-cI626 P.Stüben, det. P.Stüben, col. ZFMK CO1: GU987970 16S: GU988410


ITALY: Campania, Monti Picentini, 9 km N Acerno, Piano Laceno, 9235/I-634-cis N40°48'58", E15°07'35", Fagus, 1210m, 03.07.2008, leg. P.Stüben, DNA-JJ0573/TIS-cI634 det. P.Stüben, col. ZFMK CO1: GU987976 16S: GU988416


ITALY: Abruzzo, P.N. Majella, 11 km N Roccaraso, Bosco di S. 9256/I-655-cis Antonio, N41°56'27", E14°01'41", Pyrus/Acer, 1321m, 14.07.2008, DNA-JJ0594/TIS-cI655 CO1: GU987988 leg. P.Stüben, det. P.Stüben, col. ZFMK 16S: GU988426


ITALY: Lazio, Monti Lepini, Prati, N41°28', E13°06', 31.03.2007, leg. Brandstetter, det. Brandstetter, col. ZFMK

Acalles lemur lemur (Germar, 1824)

FRANCE: Isère, 22 km S Bourgoin, La Cote St. André, N45°24'04", 8675/I-0005-leL E05°14'04", Castanea, 469m, 08.08.2005, leg. P.Stüben, det. DNA-JJ0013/TIS-cI0005 P.Stüben, col. ZFMK CO1: GU987755 16S: GU988165

9444/I-824-cis DNA-JJ0782/TIS-cI824 CO1: GU988090 16S: GU988529




39


Collecting Data



FRANCE: Isère, 19 km S Bourgoin, N Semons, N45°26'01", E05°12'06", Quercus, Fagus, 448m, 08.08.2005, leg. P.Stüben, det. P.Stüben, col. ZFMK

8676/I-0010-leL DNA-JJ0014/TIS-cI0010 CO1: EU286446 16S: EU286281


GERMANY: RLP, Mosellus valley, 1 km N Treis-Karden, N50°11'08", E07°18'20", Quercus, Carpinus, 250m, 17.09.2005, leg. P.Stüben, det. P.Stüben, col. ZFMK

8700/M-0061-leL DNA-JJ0038/TIS-cM0061 CO1: GU987771 16S: GU988181


GERMANY: RLP, Mosellus valley, 10 km S Cochem, Bullay, N50°03'27", E07°08'51", Quercus, Carpinus, 222m, 16.09.2005, leg. P.Stüben, det. P.Stüben, col. ZFMK

8701/M-0059-leL DNA-JJ0039/TIS-cM0059 CO1: GU987772 16S: GU988182


FRANCE: Isère, 12 km SW Bourgoin, NE St. Jean de Bournay, N45°31'02", E05°10'23", Quercus, Castanea, 395m, 10.08.2005, leg. P.Stüben, det. P.Stüben, col. ZFMK

8825/I-0015-leL DNA-JJ0163/TIS-cI0015 CO1: GU987809 16S: GU988225


GERMANY: RLP, Altenahr,Teufelsloch, N50°30'36", E06°59'24", 9008/D-0422-lem Quercus, Fagus, 150m, 02.11.2007, leg. P.Stüben, det. P.Stüben, col. DNA-JJ0346/TIS-cD0422 ZFMK CO1: GU987880 16S: GU988297


FRANCE: Alpes-Maritimes, 9 km N Sospel, N Moulinet, N43°57'58", E07°24'55", Castanea sativa, 1111m, 24.12.2007, leg. P.Stüben, det. P.Stüben, col. ZFMK

9071/F-438-lem DNA-JJ0409/TIS-cF438 CO1: GU987896 16S: GU988324


GERMANY: Rheinland-Pfalz, Eppenbrunn, „Lourdesgrotte", N49°06'58", E07°33'20", Fagus, 277m, 09.05.2008, leg. P.Stüben, det. P.Stüben, col. ZFMK

9364/D-673-lem DNA-JJ0702/TIS-cD673 CO1: GU988042 16S: GU988480

Acalles maraoensis Stüben, 2001

PORTUGAL: Manteigas, N40°19'26", W07°34'20", Betula, 1531m, 9102/P-540-mar 26.01.2008, leg. J.Astrin, det. J.Astrin, col. ZFMK DNA-JJ0440/TIS-cP540 CO1: GU987909 16S: GU988337


SPAIN: Cáceres, 8 km W Guadalupe, Sierra de Guadalupe, Las Villueras Mt., N39°28'00", W05°24'22", Quercus ilex, Quercus pubescens, 1475m, 11.04.2004, leg. P.Stüben, det. P.Stüben, col. ZFMK

9182/E-581-mar DNA-JJ0520/TIS-cE581 CO1: GU987943 16S: GU988382


SPAIN: Toledo, SW Navahermosa, near Hontanar, N39°35'46", W04°30'11", Quercus ilex, 1100m, 21.04.2009, leg. J.Astrin, det. P.Stüben, col. ZFMK



PORTUGAL: Sa. Estrela, S Manteigas, N40°19'26", W07°34'20", Betula, Cytisus, Erica, 1532m, 25.04.2009, leg. J.Astrin, det. P.Stüben, col. ZFMK



PORTUGAL: Sa. Estrela, SW Manteigas, N40°18'45", W07°35'03", 9477/E-857-mar Cytisus, 1558m, 25.04.2009, leg. J.Astrin, det. P.Stüben, col. ZFMK DNA-JJ0815/TIS-cE857 CO1: GU988112 16S: GU988551 ......continued on the next page


SCHÜTTE & STÜBEN


Collecting Data



9486/E-867-mar PORTUGAL: Sa. Marão, W Vila Real, Alto do Espinho, DNA-JJ0824/TIS-cE867 N41°16'15", W07°54'17", Amelanchier, Quercus pyrenaica, Cytisus, 1145m, 30.04.2009, leg. J.Astrin, det. P.Stüben, col. ZFMK CO1: GU988118 16S: GU988557


PORTUGAL: Sa. Gerês, NE Braga, S Lindoso, N41°49'48", 9488/E-869-mar W08°12'15", Amelanchier, Erica, Castanea, 974m, 01.05.2009, leg. DNA-JJ0826/TIS-cE869 J.Astrin, det. P.Stüben, col. ZFMK CO1: GU988120 16S: GU988559


SPAIN: Burgos, SE Burgos, W slope of Trigaza Mt., N42°16'36", 9494/E-875-mar W03°15'32", Cytisus, Fagus, 1496m, 05.05.2009, leg. J.Astrin, det. DNA-JJ0832/TIS-cE875 P.Stüben, col. ZFMK CO1: GU988126 16S: GU988565

Acalles micros Dieckmann, 1982

FRANCE: Isère, 9 km N Grenoble, Mont St. Martin, N45°16'15", E05°40'44", Fagus, Fraxinus, Quercus, 813m, 12.08.2005, leg. P.Stüben, det. P.Stüben, col. ZFMK

8669/I-0017-mic DNA-JJ0007/TIS-cI0017 CO1: GU987753 16S: GU988163


FRANCE: Isère, 14 km SW Bourgoin, St. Jean de Bournay, N45°30'28", E05°09'09", Castanea, Ilex, Quercus, 406m, 07.08.2005, leg. P.Stüben, det. P.Stüben, col. ZFMK

8677/I-0003-mic DNA-JJ0015/TIS-cI0003 CO1: GU987756 16S: GU988166



8826/I-0030-mic DNA-JJ0164/TIS-cI0030 CO1: EU286493 16S: EU286329


FRANCE: Isere, 4 km E La Chapelle-de-Surieu, N45°24'18‘', E04°56'26", Fagus, Alnus, 325m, 09.04.2006, leg. P.Stüben, det. P.Stüben, col. ZFMK

8952/F-0242-mic DNA-JJ0290/TIS-cF0242 CO1: GU987845 16S: GU988263


GERMANY: BW, Kaiserstuhl, "Auf dem Eck", N48°06'32", E07°40'04", Fagus, Carpinus, Quercus, 432m, 27.05.2006, leg. P.Stüben, det. P.Stüben, col. ZFMK

8953/D-0243-mic DNA-JJ0291/TIS-cD0243 CO1: GU987846 16S: GU988264


GERMANY: Rheinland-Pfalz, 12 km W Pirmasens, Walshausen, N49°13'15", E07°26'43", Fagus, 347m, 10.05.2008, leg. P.Stüben, det. P.Stüben, col. ZFMK

9363/D-672-mic DNA-JJ0701/TIS-cD672 CO1: GU988041 16S: GU988479


FRANCE: Dép. Loire, SW Pélussin, Mont Pilat, near Crêt de l'Oeillon, N45°24'24", E04°37'06", Fagus, 1122m, 15.07.2009, leg. P.Stüben, det. P.Stüben, col. ZFMK

9564/F-929-mic DNA-JJ0902/TIS-cF929 CO1: GU988157 16S: GU988600

Acalles minutissimus (LeConte 1876)

USA: Florida, Putnam County, 5 km E Melrose, N29°42', W81°59', 11974/1763-BAN 27.02.1998, leg. H.&A.Howden, det. R.Anderson, col. ZFMK DNA-0155668049/TIS-24999 CO1: KJ767020 16S: KJ767027

Acalles misellus Boheman, 1844

FRANCE: Hautes-Pyrénées, Barbazan Debat, Bois de Rébisclou, N43°11', E00°07', 370m, 03.07.2009, leg. Brustel, det. Stüben, col. ZFMK

888/684-PST DNA-100448261/TIS-20462 CO1: KC783822 16S: n.a. ......continued on the next page



41


Collecting Data



SPAIN: Barcelona, near Vallirana, N41°22'04", E01°55'05", Silax aspera, Hedera helix, Laurus, 22.06.2008, leg. J.Astrin, det. P.Stüben, col. ZFMK

9211/E-610-mis DNA-JJ0549/TIS-cE610 CO1: GU987956 16S: GU988396


SPAIN: Barcelona, S. Montseny, Collformic, Font St. Jordi, N41°48'12", E02°20'48", Rubus, Populus, 1132m, 23.06.2008, leg. J.Astrin, det. P.Stüben, col. ZFMK



SPAIN: Barcelona, S. Montseny, Tordera valley, near St. Marçal, N41°48'01", E02°25'15", 1060m, 23.06.2008, leg. J.Astrin, det. P.Stüben, col. ZFMK



SPAIN: Barcelona, St. Esteve de Palautordera, Sta. Margarida, N41°43'07", E02°25'45", Ruscus, Hedera, Platanus, 254m, 24.06.2008, leg. J.Astrin, det. P.Stüben, col. ZFMK



NETHERLANDS: Zeeland, Walcheren, NE Oostkapelle, De Manteling (Oranjezon), N51°34'49", E03°33'45", Quercus, 15.04.2009, leg. P.Stüben, det. P.Stüben, col. ZFMK

9452/NL-832-mis DNA-JJ0790/TIS-cNL832 CO1: GU988094 16S: GU988533


SPAIN: León, NE León, N Riaño, SW Posada de Valdeón, 9491/E-872-mis N43°06'54", W04°59'33", Fagus, 1431m, 04.05.2009, leg. J.Astrin, DNA-JJ0829/TIS-cE872 det. P.Stüben, col. ZFMK CO1: GU988123 16S: GU988562


SPAIN: Palencia, N Cervera de Pisuerga, Pto. de Piedrasluengas, N43°02'57", W04°27'19", Fagus, Ilex, 1314m, 04.05.2009, leg. J.Astrin, det. P.Stüben, col. ZFMK



FRANCE: Dép. Pyrénées-Atlantique, S Sarnear Col de Lizarrieta, N43°15'46", W01°36'27", Quercus, 382m, 24.07.2009, leg. P.Stüben, det. P.Stüben, col. ZFMK

9570/F-937-mis DNA-JJ0908/TIS-cF937 CO1: GU988159 16S: GU988606


FRANCE: Dép. Pyrénées-Atlantique, W Ciboure (near coast), 9571/F-938-mis N43°22'49", W01°42'59", Quercus, 41m, 26.07.2009, leg. P.Stüben, DNA-JJ0909/TIS-cF938 det. P.Stüben, col. ZFMK CO1: GU988160 16S: GU988607


SPAIN: La Rioja, E Montenegro de Cameros, N42°06'04", W02°42'38", Fagus trees near roadside, 1109m, 01.05.2013, leg. P.Stüben & A.Schütte, det. P.Stüben, col. ZFMK



SPAIN: Castilla y León, N of Almarza, N42°01'24", W02°30'46", Ilex, Quercus, 1366m, 01.05.2013, leg. P.Stüben & A.Schütte, det. P.Stüben, col. ZFMK


Acalles monasterialis SPAIN: Salamanca, 38 km NW Béjar, Sierra de la Peña de Francia, 9062/E-505-mon Stüben, 2004 Monasterio, N40°31'01", W06°10'15", Cytisus sp., 1636m, DNA-JJ0400/TIS-cE505 10.04.2004, leg. P.Stüben, det. P.Stüben, col. ZFMK CO1: GU987891 16S: GU988319 ......continued on the next page


SCHÜTTE & STÜBEN


Collecting Data


Acalles monasterialis SPAIN: Salamanca, NW Béjar, NW Monforte de la Sierra, Peña de Stüben, 2004 Francia, near cloister, N40°31'01", W06°10'15", Cytisus, 1636m, 24.04.2009, leg. J.Astrin, det. P.Stüben, col. ZFMK

9468/E-848-mon DNA-JJ0806/TIS-cE848 CO1: GU988106 16S: GU988545

Acalles spec. nov. 4

USA: Florida, Dade County, Chekika State Rec. Area, N25°36', W80°29', hardwood hammock, 12.10.1990, leg. R.Anderson, det. R.Anderson, col. ZFMK

11978/1767-BAN DNA-0155668045/TIS-25751 CO1: KJ767022 16S: KJ767031

Acalles spec. nov. 5

USA: Florida, Dade County, Miami, Charles Deering Estate Park, N25°36', W80°18', 07.10.1990, leg. R.Anderson, det. R.Anderson, col. ZFMK

11979/1768-BAN DNA-0155668044/TIS-25791 CO1: KP065590 16S: KJ767032

Acalles papei A. & F. Solari, 1905

CROATIA: Dalmatien, 4,5 km N Drnis, Promina Mts., N43°54'20", 8852/HR-0327-pap DNA-JJ0190/TIS-cHR0327 E16°10'01", Macchia, Quercus, Carpinus, Corylus, 738m, CO1: EU286507 09.07.2007, leg. P.Stüben, det. P.Stüben, col. ZFMK 16S: EU286343


GREECE: Korfu, 18 km N Kerkyra, Loutses, Megali Grava, N39°46'38", E19°53'21", Quercus pubescens, Laurus nobilis, Asphodelus, 479m, 23.09.2007, leg. P.Stüben, det. P.Stüben, col. ZFMK

8996/KO-0369-pap DNA-JJ0334/TIS-cKO0369 CO1: GU987873 16S: GU988290


GREECE: Epirus, 25 km SE Igoumenitsa, Chrisavgi, N39°26'43", E20°32'04", Quercus ilex, Platanus, 187m, 02.10.2007, leg. P.Stüben, det. P.Stüben, col. ZFMK

9006/EP-0403-pap DNA-JJ0344/TIS-cEP0403 CO1: GU987878 16S: GU988295


GREECE: Korfu, 10 km SW Kerkyra, Kato Garouna, N39°31'14", E19°51'29", Quercus ilex, 357m, 26.09.2007, leg. P.Stüben, det. P.Stüben, col. ZFMK

9368/KO-0378-pap DNA-JJ0706/TIS-cKO0378 CO1: GU988044 16S: GU988482


GREECE: Epirus, 19 km E Igoumenitsa, Petrovitsa, N39°33'30", E20°28'12", Arbutus/Platanus, 355m, 01.10.2007, leg. P.Stüben, det. P.Stüben, col. ZFMK

9370/EP-0399-pap DNA-JJ0708/TIS-cEP0399 CO1: GU988046 16S: GU988484

Acalles parasierrae Stüben, 2002

MOROCCO: Rif, 10 km W Ketama, 10 km W Ketama, N34°57'40", 9529/E-910-prs W04°40'51", Cedrus, Prunus, 1600m, 10.05.2009, leg. P.Stüben, DNA-JJ0867/TIS-cE910 det. P.Stüben, col. ZFMK CO1: GU988146 16S: GU988586

Acalles parvulus Boheman, 1837

ITALY: Abruzzo, P.N. Majella, 11 km N Roccaraso, Bosco di S. 9555/I-654-par Antonio, N41°56'27", E14°01'41", Pyrus/Acer, 1321m, 14.07.2008, DNA-JJ0893/TIS-cI654 leg. P.Stüben, det. P.Stüben, col. ZFMK CO1: GU988151 16S: GU988594

Acalles parvulus parvulus Boheman, 1837

FRANCE: Isère, 19 km S Bourgoin, N Semons, N45°26'01", E05°12'06", Quercus, Fagus, 448m, 08.08.2005, leg. P.Stüben, det. P.Stüben, col. ZFMK

8680/I-0008-par DNA-JJ0018/TIS-cI0008 CO1: EU286448 16S: EU286283


FRANCE: Isère, 10 km SE Vienne, N45°28', E04°56', 23.06.1905, leg., det., col. ZFMK

8690/D-0069-par DNA-JJ0028/TIS-cD0069 CO1: GU987764 16S: GU988174 ......continued on the next page



43


Collecting Data



GERMANY: RLP, Mosellus valley, 1 km N Treis-Karden, N50°11'08", E07°18'20", Quercus, Carpinus, 250m, 17.09.2005, leg. P.Stüben, det. P.Stüben, col. ZFMK

8698/M-0060-par DNA-JJ0036/TIS-cM0060 CO1: GU987770 16S: GU988180


FRANCE: Isère, 14 km SW Bourgoin, St. Jean de Bournay, N45°30'28", E05°09'09", Castanea, Ilex, Quercus, 406m, 07.08.2005, leg. P.Stüben, det. P.Stüben, col. ZFMK

8708/I-0002-par DNA-JJ0046/TIS-cI0002 CO1: GU987777 16S: GU988187


FRANCE: Isere, 4 km E La Chapelle-de-Surieu, N45°24'18‘', E04°56'26", Fagus, Alnus, 325m, 09.04.2006, leg. P.Stüben, det. P.Stüben, col. ZFMK

8956/F-0248-par DNA-JJ0294/TIS-cF0248 CO1: GU987849 16S: GU988267


FRANCE: PACA, 27 km W Brignoles, Massif de la Sainte Baume, N43°19'42‘', E05°45'17", Fagus, 703m, 18.04.2006, leg. P.Stüben, det. P.Stüben, col. ZFMK



GERMANY: Rheinland-Pfalz, Mosellus valley, 5 km E Cochem, "Apolloweg", N50°08'46", E07°12'51", Quercus, 237m, 04.07.2006, leg. J.Astrin and P.Stüben, det. P.Stüben, col. ZFMK

8958/D-0250-par DNA-JJ0296/TIS-cD0250 CO1: GU987851 16S: GU988269


FRANCE: Alpes-Maritimes, 6 km NE Sospel, Col de Brouis, N43°55'28", E07°28'37", broom, Helleborus, 882m, 23.12.2007, leg. P.Stüben, det. P.Stüben, col. ZFMK



ITALY: Campania, Cilento, 6 km SE Vallo d. Lucania, M. Sacro o 9222/I-621-par Gelbison, N40°12'41", E15°19'42", Fagus, 1544m, 30.06.2008, leg. DNA-JJ0560/TIS-cI621 P.Stüben, det. P.Stüben, col. ZFMK CO1: GU987966 16S: GU988406

Acalles parvulus temperei Péricart, 1987

FRANCE: Isère, 9 km N Grenoble, Mont St. Martin, N45°16'15", E05°40'13", Quercus, 719m, 12.08.2005, leg. P.Stüben, det. P.Stüben, col. ZFMK

8681/I-0023-tem DNA-JJ0019/TIS-cI0023 CO1: EU286449 16S: EU286284



8682/I-0029-tem DNA-JJ0020/TIS-cI0029 CO1: GU987758 16S: GU988168


FRANCE: Isère, 7 km N Grenoble, Massif de la Chartreus, Umg. Quaix en Ch., N45°14'53", E05°44'31", Fagus, Ilex, 613m, 13.08.2005, leg. P.Stüben, det. P.Stüben, col. ZFMK

8827/I-0042-tem DNA-JJ0165/TIS-cI0042 CO1: GU987810 16S: GU988226


FRANCE: Les Houches, Carlaveyron, N45°55'40", E06°48'17", 2119m, 24.09.2007, leg. Brustel, det. Brustel, col. ZFMK

9431/F-804-tem DNA-JJ0769/TIS-cF804 CO1: GU988080 16S: n.a.

Acalles pilula Wollaston, 1864

SPAIN: Canary Islands, El Hierro, 7 km W La Frontera, Pista Derrabado, N27°44'29", W18°03'24", Laurus azorica, 895m, 30.12.2006, leg. P.Stüben, det. P.Stüben, col. ZFMK

8742/C-0099-pil DNA-JJ0080/TIS-cC0099 CO1: EU286461 16S: EU286296 ......continued on the next page


SCHÜTTE & STÜBEN


Collecting Data



SPAIN: Canary Islands, El Hierro, 9,5 km W La Frontera, Pista de Mencafite, N27°44'06", W18°05'08", Chamaecytisus, Pericallis, Ficus, 929m, 22.12.2006, leg. P.Stüben, det. P.Stüben, col. ZFMK



SPAIN: Canary Islands, La Palma, 2,5 km W La Galga, above Cubo 8960/E-0252-pil de la Galga, N28°45'18", W17°46'37", laurisilva, Juglans, 857m, DNA-JJ0298/TIS-cE0252 03.07.2006, leg. P.Stüben, det. P.Stüben, col. ZFMK CO1: GU987853 16S: GU988271


SPAIN: Canary Islands, El Hierro, 3 km N San Andres, Montaña de 8961/E-0253-pil la Fara, N27°47'39", W17°56'55", Sonchus hierrensis, 936m, DNA-JJ0299/TIS-cE0253 25.12.2006, leg. P.Stüben, det. P.Stüben, col. ZFMK CO1: GU987854 16S: GU988272


SPAIN: Canary Islands, La Palma, 5 km SE El Paso, Cumbre Nueva, N28°37'51", W17°49'36", laurisilva, 1415m, 28.06.2006, leg. P.Stüben, det. P.Stüben, col. ZFMK


SPAIN: Canary Islands, El Hierro, 4 km SW La Frontera, El Parque, 9127/E-0254-pil N27°44'20", W18°01'30", 739m, 24.12.2006, leg. P.Stüben, det. DNA-JJ0465/TIS-cE0254 P.Stüben, col. ZFMK CO1: GU987920 16S: GU988347


SPAIN: Canary Islands, Tenerife, E Los Silos, Teno Mts. El Tanque 9287/E-698-pil Bajo, N28°21'39", W16°46'19", 534m, 03.10.2008, leg. J.Astrin and DNA-JJ0625/TIS-cE698 P.Stüben, det. P.Stüben, col. ZFMK CO1: GU988007 16S: GU988445

Acalles ptinoides (Marsham, 1802)

GERMANY: NRW, Niederrhein, Niederrhein, W Moenchengladbach, 3 km E Niederkrüchten, N51°11'56", E06°15'38", Sphagnum, 07.06.2007, leg. P.Stüben, det. P.Stüben, col. ZFMK

8872/D-0256-pti DNA-JJ0210/TIS-cD0256 CO1: EU286522 16S: EU286359


GERMANY: NRW, N Mönchengladbach, N Mönchengladbach, Helenabrunn, N51°13'34", E06°24'00", Fagus, 60m, 15.11.2007, leg. P.Stüben, det. P.Stüben, col. ZFMK

9342/D-671-pti DNA-JJ0680/TIS-cD671 CO1: GU988036 16S: GU988473


SPAIN: Asturias, S Oviedo, SW Pto. de Pajares, Valgrande ski station, N42°58'25", W05°46'30", Betula, Erica, Ilex, Cytisus, 1649m, 03.05.2009, leg. J.Astrin, det. P.Stüben, col. ZFMK

9489/E-870-pti DNA-JJ0827/TIS-cE870 CO1: GU988121 16S: GU988560


SPAIN: Asturias, SE Oviedo, nr. Tarna, N43°06'16", W05°13'03", 9490/E-871-pti Fagus, Cytisus, 1322m, 04.05.2009, leg. J.Astrin, det. P.Stüben, col. DNA-JJ0828/TIS-cE871 ZFMK CO1: GU988122 16S: GU988561


9495/E-876-pti SPAIN: Burgos, SE Burgos, W slope of Trigaza Mt., N42°16'36", W03°15'32", Cytisus, Fagus, 1496m, 05.05.2009, leg. J.Astrin, det. DNA-JJ0833/TIS-cE876 CO1: GU988127 P.Stüben, col. ZFMK 16S: GU988566


GERMANY: Rheinland-Pfalz, W Karlsruhe, Bienwald, 9562/D-927-pti N49°00'49", E08°05'18", Quercus, 130m, 12.06.2009, leg. P.Stüben, DNA-JJ0900/TIS-cD927 det. P.Stüben, col. ZFMK CO1: GU988155 16S: GU988598





45


Collecting Data


Acalles sablensis Blatchley, 1920

USA: Florida, Monroe County, Long Key State Park: Recreation Area, N24°48', W80°48', Borrichia spp., 22.05.1990, leg. R.Anderson, det. R.Anderson, col. ZFMK

11975/1764-BAN DNA-0155668048/TIS-25001 CO1: KP065589 16S: KJ767028

Acalles sardiniaensis Stüben, 2001

ITALY: Sardinia, 21 km SO Aritzo, Barbagia Seulo, Monte Arqueri, 9017/I-460-sar N39°49'02", E09°21'59", Quercus ilex, 950m, 22.10.2004, leg. DNA-JJ0355/TIS-cI460 P.Stüben, det. P.Stüben, col. ZFMK CO1: GU987882 16S: GU988303


ITALY: E-Sardinia, NE Seui: M. Perda Liana, N39°55'13", E09°24'20", Quercus ilex, Ilex, 973m, 28.09.2010, leg. P.Stüben, det. P.Stüben, col. ZFMK


ITALY: E-Sardinia, NE Seui: M. Tonneri, N39°53'59", E09°23'24", 9657/IT1074c DNA-JJ1074/TIS-cIT1074c Quercus ilex, 926m, 28.09.2010, leg. P.Stüben, det. P.Stüben, col. CO1: KF680231 ZFMK 16S: n.a.


ITALY: E-Sardinia, E Seui: M. Arqueri, N39°49'04", E09°22'02", Quercus ilex, 925m, 29.09.2010, leg. P.Stüben, det. P.Stüben, col. ZFMK

9661/IT1078c DNA-JJ1078/TIS-cIT1078c CO1: KF680232 16S: KJ720543


ITALY: E-Sardinia, E Seui: Ulassai, N39°48'35", E09°29'48", Quercus ilex, 745m, 29.09.2010, leg. P.Stüben, det. P.Stüben, col. ZFMK

9663/IT1080c DNA-JJ1080/TIS-cIT1080c CO1: KF680233 16S: n.a.


ITALY: Sardinia, NE Aritzo: Gennargentu, M. di Iscudu, N40°01'10", E09°16'39", creek: Alnus, moss, 1511m, 30.09.2010, leg. P.Stüben, det. P.Stüben, col. ZFMK



ITALY: S-Sardinia, E Cagliari, Nationalpark M. d. Sette Fratelli: M. 9670/IT1087c Funtana-Ona, N39°17'30", E09°23'17", Quercus, Arbutus, Pistacia, DNA-JJ1087/TIS-cIT1087c CO1: KF680235 596m, 03.10.2010, leg. P.Stüben, det. P.Stüben, col. ZFMK 16S: n.a.

Acalles sarothamni Stüben, 2003

SPAIN: Castilla y León, Sierra de Gredos, 14 km NE Arenas de S. 9023/E-466-sar Pedro, Pto. del Pico, N40°19'20", W05°00'50", Cytisus sp., 1396m, DNA-JJ0361/TIS-cE466 23.04.2003, leg. P.Stüben, det. P.Stüben, col. ZFMK CO1: GU987885 16S: GU988307

Acalles sarothamni Stüben, 2003

SPAIN: Ávila, 46 km SW Ávila, Sierra de Villafranca, Navacepedilla, N40°28'59", W05°11'08", Cytisus, 1266m, 06.04.2004, leg. P.Stüben, det. P.Stüben, col. ZFMK

9024/E-467-sar DNA-JJ0362/TIS-cE467 CO1: GU987886 16S: GU988308

Acalles sierrae H. Brisout de Barneville, 1865

SPAIN: Andalucía, 12 km N Laujar, Sierra Nevada, N37°05'45", W02°57'37", Erinacea anthyllis, 2285m, 15.10.2005, leg. P.Stüben, det. P.Stüben, col. ZFMK

8810/E-0100-sie DNA-JJ0148/TIS-cE0100 CO1: EU286492 16S: EU286328


SPAIN: Andalucía, 17 km NW Laujar, Sierra Nevada, S Puerto de la 8968/E-0261-sie Ragua, N37°06'11", W03°01'40", Bupleurum, broom, 2070m, DNA-JJ0306/TIS-cE0261 12.10.2005, leg. P.Stüben, det. P.Stüben, col. ZFMK CO1: GU987859 16S: GU988277




SCHÜTTE & STÜBEN


Collecting Data



SPAIN: Granada, NE of Granada: Sierra de La Yedra, N37°15'50", W03°32'13", Quercus ilex, 1414m, 13.05.2013, leg. P.Stüben & A.Schütte, det. P.Stüben, col. ZFMK



SPAIN: Jaen, SE of Mancha Real: Sierra Almaden (El Almaden, Locus typicus), N37°44'09", W03°31'26", Euphorbia nicaeensis, Bupleurum spinosum, 1963m, 15.05.2013, leg. P.Stüben & A.Schütte, det. P.Stüben, col. ZFMK



SPAIN: Jaen, SE of Mancha Real: Sierra Almaden (El Almaden), N37°44'23", W03°32'07", Erinacea anthyllis, 1662m, 15.05.2013, leg. P.Stüben & A.Schütte, det. P.Stüben, col. ZFMK

10210/1310-PST DNA-100426166/TIS-4126 CO1: KF680249 16S: n.a.


SPAIN: Jaen, Sierra Magina: SE of Torres, N37°44'46", W03°29'53", Erinacea anthyllis, Quercus ilex., Astragalus spec., Bupleurum, 1527m, 19.05.2013, leg. P.Stüben & A.Schütte, det. P.Stüben, col. ZFMK



SPAIN: Jaen, Sierra Magina: SE of Torres, N37°44'06", W03°30'13", Erinacea anthyllis, Euphorbia nicaeensis, 1663m, 19.05.2013, leg. P.Stüben & A.Schütte, det. P.Stüben, col. ZFMK



SPAIN: Jaen, S of La Guardia: Sierra de la Pandera, N37°40'20", 10244/1344-PST W03°41'48", Erinacea anthyllis, Cytisus spec., 1223m, 19.05.2013, DNA-100426265/TIS-4160 leg. P.Stüben & A.Schütte, det. P.Stüben, col. ZFMK CO1: KF680255 16S: KJ720562

Acalles sierrae SPAIN: Granada, E of Baul, Sierra de Baza: Santa Barbara, (formerly: bazaensis) N37°22'25", W02°51'03", Bupleurum, (Astragalus spec), 2023m, Stüben, 2001 17.05.2013, leg. P.Stüben & A.Schütte, det. P.Stüben, col. ZFMK


SPAIN: Granada, E of Baul, Sierra de Baza: Santa Barbara, Acalles sierrae (formerly: bazaensis) N37°22'50", W02°51'06", Bupleurum, (Astragalus spec), 1990m, 17.05.2013, leg. P.Stüben & A.Schütte, det. P.Stüben, col. ZFMK Stüben, 2001


Acalles sintraniensis Stüben, 1999

PORTUGAL: Sa. Estrela, S Manteigas, N40°19'26", W07°34'20", Betula, Cytisus, Erica, 1532m, 25.04.2009, leg. J.Astrin, det. P.Stüben, col. ZFMK

9473/E-853-sin DNA-JJ0811/TIS-cE853 CO1: GU988110 16S: GU988549


PORTUGAL: Sa. Estrela, SW Manteigas, N40°18'45", W07°35'03", 9476/E-856-sin Cytisus, 1558m, 25.04.2009, leg. J.Astrin, det. P.Stüben, col. ZFMK DNA-JJ0814/TIS-cE856 CO1: GU988111 16S: GU988550


PORTUGAL: Minho, NW Ponte de Lima, Vila Nova de Cerveira, Rio Coura, N41°52'27", W08°41'57", 123m, 07.05.2012, leg. P.Stüben, det. P.Stüben, col. ZFMK



PORTUGAL: Minho, PN Peneda-Geres, near Soajo, N41°52'14", W08°16'30", Cytisus, 200m, 09.05.2012, leg. P.Stüben, det. P.Stüben, col. ZFMK

9734/833-PST DNA-100414258/TIS-3746 CO1: KF680239 16S: KJ720549 ......continued on the next page



47


Collecting Data


Acalles spec.

MEXICO: Chiapas, Reserva Huitepec, leaf litter under pines, 2600m, 11.07.2007, leg. J.Longino with R.Anderson, det. J. Longino with R. Anderson, col. ZFMK

8883/X-0361-AcS DNA-JJ0221/TIS-cX0361 CO1: EU286526 16S: EU286363

Acalles sylvosus Blatchley, 1916

USA: Florida, Dade County, Miami, Camp Mahachee, N25°40', W80°16', hardwood hammock, 17.10.1990, leg. R.Anderson, det. R.Anderson, col. ZFMK

11976/1765-BAN DNA-0155668047/TIS-25739 CO1: KJ767021 16S: KJ767029

Acalles testensis Stüben, 2003

MOROCCO: High Atlas, N Tizi-n-Test, Tachguette, N30°52'25", W08°21'25", Euphorbia nicaeensis, Quercus ilex, broom, 2047m, 30.04.2009, leg. P.Stüben, det. P.Stüben, col. ZFMK

9499/E-880-tes DNA-JJ0837/TIS-cE880 CO1: GU988128 16S: GU988568

Acalles tibialis (Weise, 1891)

FRANCE: Alpes-Maritimes, 8 km NW Sospel (2 km S Peira Cava), 9068/F-435-tib DNA-JJ0406/TIS-cF435 Baisse de la Cabanette, N43°54'53", E07°21'11", Fagus, 1371m, CO1: GU987895 24.12.2007, leg. P.Stüben, det. P.Stüben, col. ZFMK 16S: GU988323

Acalles tibialis (Weise, 1891)

FRANCE: Alpes-Maritimes, 3 km W Sospel, Col de Braus, 9075/F-442-tib N43°52'34", E07°24'17", Quercus pubescens, Ostrya carpinifolia, DNA-JJ0413/TIS-cF442 broom, 1051m, 26.12.2007, leg. P.Stüben, det. P.Stüben, col. ZFMK CO1: GU987898 16S: GU988326

Acalles vorsti spec.nov. Stüben, 2014

SPAIN: Mallorca, Soller: Son Coll, N39°46'06", E02°40'12", Vorst, 10258/1358-PST Quercus forest, 287m, 04.04.2002, leg. O.Vorst, det. P.Stüben, col. DNA-100417602/TIS-22183 ZFMK CO1: KF680256 16S: KJ720563

Acallocrates denticollis (Germar, 1824)

CROATIA: Dalmatien, 4,5 km N Drnis, Promina Mts., N43°54'20", 8988/HR-0325-den E16°10'01", Quercus, Carpinus, Corylus, 738m, 09.07.2007, leg. DNA-JJ0326/TIS-cHR0325 P.Stüben, det. P.Stüben, col. ZFMK CO1: GU987868 16S: GU988285

Acallocrates minutesquamosus (Reiche, 1860)

FRANCE: PACA, 27 km W Brignoles, Massif de la Sainte Baume, N43°19'42", E05°45'17", Fagus, 703m, 18.04.2006, leg. P.Stüben, det. P.Stüben, col. ZFMK

Acallorneuma doderoi A. & F. Solari, 1909

ITALY: Sicilia (PA), 6 km SW Godrano, Bosco Ficuzza, Mte. Rocca 8727/S-0082-dod Busambra, N37°51'38", E13°23'24", Quercus, Fraxinus, 1200m, DNA-JJ0065/TIS-cS0082 16.10.2002, leg. P.Stüben, det. P.Stüben, col. ZFMK CO1: EU286457 16S: EU286292

Acallorneuma ITALY: E-Sardinia, N Dorgali: M. Tuttavista (Gipfel), N40°22'46", sardiniense E09°38'22", Quercus ilex, 801m, 06.10.2010, leg. P.Stüben, det. Osella & Zuppa, 2002 P.Stüben, col. ZFMK

8847/F-0262-min DNA-JJ0185/TIS-cF0262 CO1: EU286504 16S: EU286340

9679/IT1096c DNA-JJ1096/TIS-cIT1096c CO1: JX181781 16S: n.a.

Aeoniacalles aeonii aeonii (Wollaston, 1864)

8811/C-0101-aeo SPAIN: Canary Islands, Tenerife, Anaga Mts., 10 km NE La Laguna, Afur, N28°33'10", W16°14'59", Aeonium cuneatum, 300m, DNA-JJ0149/TIS-cC0101 CO1: GU987800 29.12.2003, leg. P.Stüben, det. P.Stüben, col. ZFMK 16S: GU988217

Aeoniacalles aeonii bodegensis (Wollaston, 1864)

SPAIN: Canary Islands, Tenerife, East Anaga Mts., 19 km NE La Laguna, Lomo de las Bodegas, N28°33'43", W16°09'25", Aeonium holochrysum, 500m, 22.12.2003, leg. P.Stüben, det. P.Stüben, col. ZFMK

9148/E-547-bod DNA-JJ0486/TIS-cE547 CO1: GU987927 16S: GU988358 ......continued on the next page


SCHÜTTE & STÜBEN


Collecting Data


Calacalles bandamaensis Stüben, 2012

SPAIN: Canary Islands, Gran Canaria, Marzagan: Caldera de Bandama, N28°02'07", W15°27'36", Ficus carica, 401m, 23.12.2010, leg. P.Stüben, det. P.Stüben, col. ZFMK

197/7-PST_2GC2011 DNA-100438112/TIS2D100446901 CO1: JN701880 16S: n.a.

Calacalles nataliae Astrin & Stüben, 2009

SPAIN: Canary Islands, La Gomera, SE Hermigua, near Casas del Palmar, N28°09'29", W17°09'37", thermophilous shrub forest, 627m, 08.10.2008, leg. J.Astrin and P.Stüben, det. P.Stüben, col. ZFMK

9345/E-744-nat DNA-JJ0683/TIS-cE744 CO1: GU988037 16S: GU988474

Canariacalles alluaudi (Uyttenboogaart, 1940)

SPAIN: Canary Islands, Tenerife, W Los Silos, Teno Mts. Casa Blanca near Buenavista del Norte, N28°21'36", W16°52'10", Ceropegia dichotoma, 231m, 02.10.2008, leg. J.Astrin and P.Stüben, det. P.Stüben, col. ZFMK

9284/E-695-all DNA-JJ0622/TIS-cE695 CO1: GQ332330 16S: GQ332314

Coloracalles edoughensis comb. nov. (Desbrochers, 1892)

SPAIN: Barcelona, near Vallirana, N41°22'36", E01°55'02", Quercus ilex, Ficus carica, Smilax aspera, 245m, 22.06.2008, leg. J.Astrin, det. P.Stüben, col. ZFMK

9212/E-611-edo DNA-JJ0550/TIS-cE611 CO1: GU987957 16S: GU988397

Coloracalles humerosus (Fairmaire, 1862)

SPAIN: Castellón, Morella, Barranco de la Bota, N40°33'12", W00°00'27", Quercus ilex, Hedera helix, 814m, 21.06.2008, leg. J.Astrin, det. P.Stüben, col. ZFMK

9207/E-606-hum DNA-JJ0545/TIS-cE606 CO1: GU987952 16S: GU988392

Cryptorhynchus lapathi (Linné, 1758)

GERMANY: Bienen bei Rees, old arm of Rhine river, N51°46', E06°20', 19.05.2004, leg. S.Scharf, det. S.Scharf, col. ZFMK

8876/D-0354-lap DNA-JJ0214/TIS-cD0354 CO1: EU286523 16S: EU286360

Dendroacalles fortunatus (Wollaston, 1864)

SPAIN: Canary Islands, La Gomera, SW Hermigua, Los Aceviños, N28°08'24", W17°13'45", Persea indica, 992m, 11.10.2008, leg. Astrin, Stüben, Behne and Floren, det. P.Stüben, col. ZFMK

9435/E-808-for DNA-JJ0773/TIS-cE808 CO1: GU988082 16S: GU988521

Dendroacalles ornatus (Wollaston, 1854)

PORTUGAL: Madeira, 3 km W Seixal, "Lagoa" near "Fanal", N32°48'47", W17°08'59", Ocotea foetens, 950m, 22.03.2008, leg. J.Astrin and P.Stüben, det. J.Astrin and P.Stüben, col. ZFMK

9094/P-530-orn DNA-JJ0432/TIS-cP530 CO1: FJ716548 16S: FJ716497

Dichromacalles diocletianus (Germar, 1817)

FRANCE: Dép. Vaucluse, Avignon, near Montfavet, N43°56'20", E04°50'47", Picris echioides, 31m, 19.07.2009, leg. P.Stüben, det. P.Stüben, col. ZFMK

9557/F-933-dio DNA-JJ0895/TIS-cF933 CO1: GU988152 16S: GU988595

Dichromacalles dromedarius (Boheman, 1844)

9113/P-515-dro PORTUGAL: Madeira, 1 km NE Machico, Pico do Facho, N32°43'33", W16°45'34", Tolpis succulenta, 212m, 17.03.2008, leg. DNA-JJ0451/TIS-cP515 CO1: GU987917 P.Stüben, det. P.Stüben, col. ZFMK 16S: GU988344

Echinodera crenata Wollaston, 1863

SPAIN: Canary Islands, Tenerife, S Orotava, Orotava valley, Mirador de la Rosa, N28°20'25", W16°31'29", Greenovia sp., Aeonium spathulatum, 1503m, 04.10.2008, leg. J.Astrin and P.Stüben, det. P.Stüben, col. ZFMK

Echinodera hypocrita FRANCE: Isère, 12 km SW Bourgoin, NE St. Jean de Bournay, (Boheman, 1837) N45°31'02", E05°10'23", Quercus, Castanea, 395m, 10.08.2005, leg. P.Stüben, det. P.Stüben, col. ZFMK

9299/E-710-cre DNA-JJ0637/TIS-cE710 CO1: GU213719 16S: GU213584 8831/I-0014-hyp DNA-JJ0169/TIS-cI0014 CO1: GU213647 16S: GU213512 ......continued on the next page



49


Collecting Data


Elliptacalles baeticus SPAIN: Málaga, W Estepona, N36°25'58", W05°14'42", Quercus, Stüben, 2008 Ceratonia, Pistacia, 189m, 13.01.2009, leg. Stüben, det. P.Stüben, col. ZFMK

9401/E-768-bae DNA-JJ0739/TIS-cE768 CO1: GU988060 16S: GU988500

Elliptacalles longus (Desbrochers, 1892)

ITALY: Sicilia (PA), 6 km SW Carini, Montagna Longa, N38°07'10", E13°08'31", Quercus ilex, 647m, 09.10.2006, leg. P.Stüben, det. P.Stüben, col. ZFMK

8739/I-0096-lon DNA-JJ0077/TIS-cI0096 CO1: EU286458 16S: EU286293

Eurhoptus spec.

MEXICO: Chiapas, Reserva Huitepec, leaf litter under pines, 2600m, 11.07.2007, leg. J.Longino with R.Anderson, det. R. Anderson, col. ZFMK

8881/X-0359-EuS DNA-JJ0219/TIS-cX0359 CO1: EU286524 16S: EU286361

Ficusacalles senilis senilis (Wollaston, 1864)

8732/C-0087-sen SPAIN: Canary Islands, El Hierro, 3 km N San Andres, N slope Montaña de la Fara, N27°47'39", W17°56'55", Foeniculum vulgare, DNA-JJ0070/TIS-cC0087 CO1: GU987784 936m, 25.12.2006, leg. P.Stüben, det. P.Stüben, col. ZFMK 16S: GU988198

Gasterocercus depressirostris depressirostris (Fabricius, 1792)

GERMANY: Rhineland-Palatinate, "Biehnwald" conservation area, Collector´s No: leg. F.Köhler BFB_Col_FK_2246 CO1: will be released later by GBOL Munich

Kyklioacalles plantapilosus Stüben & Astrin, 2010

SPAIN: Almería, 11 km NW Laujar de Andarax, Sierra Nevada, Bayárcal, N37°02'27", W03°00'12", Euphorbia, 1291m, 07.04.2007, leg. J.Astrin and P.Stüben, det. P.Stüben, col. ZFMK

8887/E-0190-pla DNA-JJ0225/TIS-cE0190 CO1: GU981475 16S: GU981523

Kyklioacalles roboris GERMANY: RLP, Moselle valley, 6 km SE Cochem, above Ediger, 8703/M-0054-rob (Curtis, 1834) N50°05'28", E07°11'01", Quercus, Carpinus, 270m, 16.09.2005, DNA-JJ0041/TIS-cM0054 leg. P.Stüben, det. P.Stüben, col. ZFMK CO1: GU987773 16S: GU988183 Kyklioacalles solarii (Fiori, 1903)

ITALY: Toscana, Passo del Spino, 3 km E Chiusi della Verna (AR), 9030/I-473-sol DNA-JJ0368/TIS-cI473 N43°42'08", E11°57'13", Fraxinus, 1000m, 18.10.2001, leg. CO1: GU981486 P.Stüben, det. P.Stüben, col. ZFMK 16S: GU981534

Lauriacalles acutus (Wollaston, 1864)

SPAIN: Canary Islands, La Palma, 2,5 km W La Galga, above Cubo 8774/C-0137-act de la Galga, N28°45'18", W17°46'37", Laurisilva, 857m, DNA-JJ0112/TIS-cC0137 03.07.2006, leg. P.Stüben, det. P.Stüben, col. ZFMK CO1: FJ716538 16S: FJ716486

Madeiracalles dispar PORTUGAL: Madeira, 3 km NE Prazeres, "Achada Grande", (Wollaston, 1854) N32°47'05", W17°11'06", Laurisilva, 1189m, 23.03.2008, leg. J.Astrin and P.Stüben, det. J.Astrin and P.Stüben, col. ZFMK

9095/P-531-dis DNA-JJ0433/TIS-cP531 CO1: FJ716549 16S: FJ716498

Madeiracalles machadoi (Stüben, 2006)

PORTUGAL: Madeira, 6 km N Ponta do Sol, Paul da Serra, "Loiral", N32°44'20", W17°06'09", Cytisus scoparius, 1295m, 18.03.2008, leg. J.Astrin and P.Stüben, det. J.Astrin and P.Stüben, col. ZFMK

9115/P-517-mac DNA-JJ0453/TIS-cP517 CO1: FJ716556 16S: FJ716506

Montanacalles nevadaensis (Stüben, 2001)

SPAIN: Almería, 11 km NW Laujar de Andarax, Sierra Nevada, Bayárcal, N37°02'27", W03°00'12", Quercus ilex, broom, Euphorbia, 1291m, 07.04.2007, leg. J.Astrin and P.Stüben, det. P.Stüben, col. ZFMK

8837/E-0189-nev DNA-JJ0175/TIS-cE0189 CO1: EU286498 16S: EU286334 ......continued on the next page


SCHÜTTE & STÜBEN


Collecting Data


Onyxacalles balearicus Stüben, 2005

SPAIN: Mallorca, 3 km SE Lluc, Sra. de Tramuntana, Sa Maleta, N39°48'47", E02°53'23", Quercus ilex, 571m, 22.08.2004, leg. P.Stüben, det. P.Stüben, col. ZFMK

8869/E-0168-bal DNA-JJ0207/TIS-cE0168 CO1: EU286521 16S: EU286357

Onyxacalles luigionii ITALY: Campania, Monti Picentini, 9 km N Acerno, Piano Laceno, 9236/I-635-lui (A. & F. Solari, 1907) N40°48'58", E15°07'35", Fagus, 1210m, 03.07.2008, leg. P.Stüben, DNA-JJ0574/TIS-cI635 det. P.Stüben, col. ZFMK CO1: GU987977 16S: GU988417 Paratorneuma aphroditae Germann & Stüben, 2006

SPAIN: Canary Islands, La Gomera: S Hermigua, El Cedro: Meriga, N28°09'12", W17°14'13", Laurisilva (river valley): Persea indica (soil sample from locus tyicus), 829m, 28.12.2009, leg. P.Stüben, det. P.Stüben, col. ZFMK

9597/ES1014 DNA-JJ1014/TIS-cES1014 CO1: JX181783 16S: JX181782

Paratorneuma orbatum (Wollaston, 1865)

SPAIN: Canary Islands, La Gomera, Hermigua, Ibo Alfaro, N28°09'58", W17°12'11", Persea, 255m, 09.10.2008, leg. J.Astrin and P.Stüben, det. P.Stüben, col. ZFMK

9322/E-733-orb DNA-JJ0660/TIS-cE733 CO1: FJ716580 16S: FJ716532

Pseudodichromacalle SPAIN: Canary Islands, Tenerife, NE La Laguna, Anaga Mts. Las s fernandezi Bodegas, N28°33'42", W16°09'25", Sonchus acaulis, 502m, (Roudier, 1954) 30.09.2008, leg. J.Astrin and P.Stüben, det. P.Stüben, col. ZFMK

9276/E-687-fer DNA-JJ0614/TIS-cE687 CO1: GU988002 16S: GU988440

Silvacalles instabilis (Wollaston, 1864)

SPAIN: Canary Islands, Tenerife, NE La Laguna, Anaga Mts. near Chinobre, N28°33'21", W16°10'46", Laurus, Ixanthus viscosus, 808m, 30.09.2008, leg. J.Astrin and P.Stüben, det. P.Stüben, col. ZFMK

9273/E-684-ins DNA-JJ0611/TIS-cE684 CO1: GU988000 16S: GU988438

Silvacalles lunulatus (Wollaston, 1854)

PORTUGAL: Madeira, 4,5 km S São VicentBoca da Encumeada, "Folhadal", N32°45'08", W17°01'40", Laurisilva, Euphorbia mellifera, 1004m, 22.03.2008, leg. J.Astrin and P.Stüben, det. J.Astrin and P.Stüben, col. ZFMK

9091/P-527-lun DNA-JJ0429/TIS-cP527 CO1: GU987906 16S: GU988334

Sonchiacalles muelleri (Stüben, 2000)

SPAIN: Canary Islands, La Palma, 4,5 km E Puntagorda, Lomo de la Ciudad, N28°46'24", W17°56'16", Tolpis lagopoda, 1241m, 15.07.2006, leg. P.Stüben, det. P.Stüben, col. ZFMK

8955/E-0246-mue DNA-JJ0293/TIS-cE0246 CO1: GU987848 16S: GU988266

Sonchiacalles sonchi (Stüben, 2000)

SPAIN: Canary Islands, Tenerife, W Los Silos, Teno Mts. Casa 9286/E-697-son Blanca near Buenavista del Norte, N28°21'36", W16°52'10", 231m, DNA-JJ0624/TIS-cE697 02.10.2008, leg. J.Astrin and P.Stüben, det. P.Stüben, col. ZFMK CO1: GU988006 16S: GU988444

Torneuma caecum Wollaston, 1860

PORTUGAL: Madeira, 1 km S Curral das Freiras, Seara Velha, N32°42'35", W16°58'17", Ficus carica, 384m, 20.03.2008, leg. J.Astrin and P.Stüben, det. J.Astrin and P.Stüben, col. ZFMK

9120/P-523-cae DNA-JJ0458/TIS-cP523 CO1: FJ716559 16S: FJ716509

Tylodinus spec.

MEXICO: Chiapas, Reserva Huitepec, leaf litter under pines, 2600m, 11.07.2007, leg. J.Longino with R.Anderson, det. R. Anderson, col. ZFMK

8882/X-0360-TyS DNA-JJ0220/TIS-cX0360 CO1: EU286525 16S: EU286362



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