Miscel.lania Zoologica 21.1 (1998) 75

The family MacLeay, 1825 (Coleoptera, Dryopoidea) in the lberian peninsula and the Balearic lslands

P. Aguilera, A. Mascagni & l. Ribera

Aguilera, P., Mascagni, A. & Ribera, l., 1998. The family Heteroceridae MacLeay, 1825 (Coleoptera, Dryopoidea) in the lberian peninsula and the Balearic Islands. Misc. Zool., 21 .l:75-100.

The family Heteroceridae MacLeay, 1825 (Coleoptera, Dryopoidea) in the lberian peninsula and the Balearic 1slands.-The information available on the family Heteroceridae in the lberian peninsula and the Balearic lslands is revised, based on published sources and on the study of unpublished material from the authors and museum collections. The male genitalia of the 17 studied species, eight Augyles and nine , is illustrated, and a brief description is included. Augyles hispidulus Kiesenwetter is recorded for the first time in the lberian peninsula, in Spain, and Heterocerus fossor Kiesenwetter for the first time in Portugal. A key to the European genus and the lberian and Balearic species of Heteroceridae is provided. Brief ecological notes and distribution maps of al1 species are also included. While the known number of species in Spain, eight Augyles and eight Heterocerus, is similar to other nearby geographical areas, the fauna of Portugal, with only seven species, and that of the Balearic Islands, with three, is considered to be still poorly known. The possible unrecorded presence of central European and north African species in Spain is discussed.

Key words: Coleoptera, Heteroceridae, Distribution, lberian peninsula, Balearic Islands.

(Rebut: 15 XII 97; Acceptació condicional: 28 IV 98; Acc. definitiva: 9 VI 98)

Pedro Aguilera, Museu de Zoologia, Ap. de correus 593, 08080 Barcelona, Espanya (Spain).- Ignacio Ribera, Dept. of Entomology, The Natural History Museum, London SW7 SBD, UK.- Alessandro Mascagni, Museo di Storia Naturale "La Specola", Universita degli Studi di Firenze, Via Romana 17, 50 125 Firenze, Italia (Italy).

ISSN: 021 1-6529 O 1998 Museu de Zoologia 76 Aauilera et al. lntroduction For more details on the biology and the mor- phology of the species of the family see CLARKE According to CROWSON(1960) the family (1973) or MASCAGNI(in press). Heteroceridae is included in Dryopoidea, to- lnformation on the family Heteroceridae gether with Dryopidae, Elmidae, Limnichidae, in the lberian peninsula is scarce, with only and Lutrochidae (the latter present only in isolated references in general works, most the American continent). This is the system- previous to the catalogue by FUENTE (1929). atic arrangement with the widest acceptance The only recent records are in more general at the moment (see e.g. MASCAGNI,in press), papers about the aquatic Coleoptera of although LAWRENCE& NEVVTON (1995) have re- Aragón (RIBERAet al., 1996). Catalonia (RIBERA cently proposed a new ordination, including & AGUILERA,1996), or papers including records al1 families of Dryopoidea, together with from the lberian peninsula and the Balearic Psephenoidea and family Byrrhidae, in the lslands (MASCAGNI,1985a. now partially up- same superfamily . According to dated, and MASCAGNI,1991). A revision of the these authors, the family is divided in two of the European species of subfamilies: Elythomerinae Pacheco, 1964, ex- Heteroceridae is about to be published clusively from Australia, and Heteroceririae (MASCAGNI,in press) but it does not include MacLeay, 1825, with an almost cosmopolitan detailed information on the distribution of distribution (LAWRENCE& NEWON, 1995). In this the lberian species. paper the classification used by MASCAGNI(in This paper gathers the information avail- press), in which Heteroceridae are separated able on this group of Coleoptera in the in five tribes (Micilini, Augyliini, Heterocerini, lberian peninsula and the Balearic Islands, Tropicini and Elythomerini), is followed. including al1 published references together Three tribes of Heterocerinae are found with unpublished information from muse- in Europe, with three genera: Augyles ums and private collections. The aedeagus (Augyliini), Heterocerus (Heterocerini) and of al1 species is illustrated, and a brief de- Micílus (Micilini) (the latter apparently not scription is provided, together with a classi- present in the lberian peninsula). All three fication key (for a more complete descrip- (and in general the whole family) have a tion of the morphology of al1 European spe- very similar morphology, with a small to me- cies see MASCAGNI,in press). dium size (approximately from 1.3 to 7.6 mm), and an elongated, parallel-sided body, cov- ered with dense pubescence. The antennae Material and methods have between nine and 11 segments, with the last six or seven forming an elongated Because of their occurrence on the interface club (except in Micilus, in which only the between the aquatic and the terrestrial envi- last segment is enlarged). The mandibles are ronments, heterocerids are frequently ignored prominent, and the anterior legs are modi- by specialists of both faunas. They are only fied for digging: they are robust, with nu- occasionally captured by usual methods for merous spines, and with an enlarged tibia. collecting aquatic Coleoptera, and when this All species have a series of stridulatory striae happens they are frequently overlooked due at both sides of the first abdominal segment, to the extraordinary velocity with which they with a different arrangement depending on may fly. There are two basic methods to collect the genus. heterocerids: to search visually on the banks The species of Heteroceridae are semi- and the shores of the water bodies, or to aquatic, and are usually found in sandy or "wash" these banks and shores, collecting the muddy shores or river banks, in which they washed material on the water surface with a dig colonial galleries. More than one species net. Water poured over the shore floods the may coexist in the same habitat. They can also galleries in which heterocerids live, forcing be found in inter-tidal marshes, forming colo- the to emerge. They are then swept to nies, in the sand. They do not occur habitii- the water surface. Due to the dense hydro- ally in mountain areas, in very stony shores, or phobic pubescence covering their entire bod- in areas frequented by cattle (probably due ies the heteroceridsfloat on the surface of the to the mechanical perturbation of the soil). water, from where they can take off directly. Miscel.lania Zoologica 21.1 (1 998)

The distributional areas of the species were markings; copious pubescence interposedwith defined according to the typology proposed longer setae; coarsely and densely punc- by VIGNATAGLIANTI et al. (1992). The records of turated; striations faint. Legs ochre with whit- "Pirineos orientales" (eastern Pyrenees) and ish setae; tibiae with nurnerous spines. Abdo- "Pirineos occidentales" (western Pyrenees) by men black or dark brown with ochre border; FUENTE(1929) have been included, although stridulatory ridge rnarked, striae evident; post- rnost probably these refer to the north side, rnetacoxal ridge complete; copious whitish in France. Descriptions of species, figures of pubescence interposed with longer setae. Sirni- aedeagus, and keys follow MASCAGNI(in press). lar species: A. hispidulus (Kiesenwetter) and Acronyrns used in the text: CARB. Coll. P. A. pruínosus (Kiesenwetter). Aguilera & l. Ribera, Barcelona; CFA. Coll. G. N. Foster, Ayr; CHB. Coll. C. Hernando, Bar- Distribution celona; CMM. Coll. A. Millán, Murcia; CMS. Western Mediterranean. Coll. A. Mascagni, Scandicci; CSCR. Coll. European distribution: western Mediterra- Skalicky, Ceska Republic; MCG. Museo Civico nean (WINKLER,1924); Spain, southern France, di Storia Naturale di Genova; MCM. Museo Morocco (ZAITZEV,1924); France (BARTHE,1926); Civico di Storia Naturale di Milano; MCR. ltaly (MASCAGNI& CALAMANDREI, 1992). Museo Civico di Zoologia, Roma; MCT. lberian records (fig. 18): Spain. Spain Museo Civico di Storia Naturale di Trieste; (BARTHE,1926); Sevilla: Castillejo, Guadajoz MCVN. Museo Civico di Storia Naturale di (KIESENWETER,1865); Andalucía (KUWERT,1890); Venezia; MCVR. Museo Civico di Storia Natu- Cádiz: salinas (ROSENHAUER,1856); Badajoz, rale di Verona; MNB. Museurn für Natur- Ciudad Real, Madrid, Sevilla (FUENTE,1929). kunde, Universitat .Berlin; MNCN. Museo Portugal. Estrernadura: Azarnbuja (PAULINODE Nacional de Ciencias Naturales, Madrid; MZB. OLIVEIRA,1882, as curtulus; FUENTE, 1929; Museu de Zoologia, Barcelona; MZF. Museo MASCAGNI,1992). Zoologico della Universita di Fiorenze; NMW. Naturhistorisches Museurn Wien. Studied material (fig. 18) Spain. Cádiz: 3 exs. 4V 1994, Algeciras, leg. W. Liedrnan (MCM), 1 ex. sarne (CMS). Results Castellón: 1 ex., Fuente la Reina (MNCN). Madrid: 3 exs. (MNCN); 1 ex., coll. Pérez Tribu Augyliini Arcas (MNCN). Málaga: 1 ex. 5 V 1991, Si- erra de Ronda (Puerto del Viento 1.100- Augyles Schiodte, 1866 1.300 m.), leg. F. Hieke (MNB). Oviedo: 1 ex. Subgenus Littorimus Gozis, 1885 10 IV 1929 (MNCN); 2 exs., Villaviciosa (MNCN). Sevilla: 1 ex., leg. Snizek (CSK); Augyles (Littorimus) curtus (Rosenhauer) 1 ex. IV 1917 (MNCN); 2 exs. 11 V 1991, Heterocerus curtus Rosenhauer,Thiere Andal. 1856: 116 Lebrija, leg. Snizek (CSK), 1 ex. sarne (CMS). gravidus Kiesenwetter, 1850 Portugal. Algarve: 1 ex. V 1910, Mon- curtulus Fairmaire, 1868 chique, leg. Schatzrnayr (CMS). Alto Alentejo: curtus curtinigripes Kuwert, 1890 3 exs. V 1910, Évora (MCG), 1 ex. idern (CMS), andalusiacus Breit, 1916 1 ex. idern (MCM); Estrernadura: 1 ex., Queluz, leg. Schatzmayr (MCT). Aedeagus as in figure 1. Total length 3.0- 3.5 mrn. Head black, whitish pubescence with According to BARTHE(1926) and MASCAGNI(1992) longer setae; mandibles dentate, rather short; this species is found on sandy substraturn. clypeus with fine copious setae, antennae with 11 segrnents and 7-segrnented club. Pronoturn black or dark brown sornetirnes with light Augyles (Littorimus) flavidus (Rossi) brown border and middle line; narrowing Heterocerus flavidus Rossi, Mant. Ins. Etr. 11, 1794: 79 anteriorly and bordered posteriorly; whitish minutus Kiesenwetter, 1835 pubescence with longer setae especially on minimus Kiesenwetter, 1843 the sides; coarsely and densely puncturated. campestris Motschulsky, 1853 Elytra brown or reddish brown with ochre flavescens Schaufuss, 1862 78 Aguilera et al. pallescens Schaufuss, 1862 though MASCAGN~(1992) also recorded it albipennis Kuwert, 1890 from muddy substratum.

Aedeagus as in figure 2. Total length 2.0- 3.0 mm. Head brown, whitish pubescence Augyles (Littorimus) hispidulus (Kiesenwetter) with some longer setae especially on the Heterocerus hispidulus Kiesenwetter, Germ. sides; mandibles dentate with an acute apex; Zeitschr. IV, 1843: 21 1, t. 3, f. 7 clypeus with fine copious setae; antennae with 10 segments and 6-segmented club. Aedeagus as in figure 6. Total length 2.9- Pronotum brown with light brown border 3.6 mm. Head black, whitish pubescence inter- and occasionally a light-brown middle line; posed with longer setae; mandibles dentate, narrowing anteriorly and bordered post- rather short, with rounded apex; clypeus with eriorly; whitish pubescence with some longer fine copious setae; antennae with 10 segments setae especially on the sides; finely and and 6-segrnented club. Pronotum black, some- densely puncturated. Elytra light brown with times with reddish border and middle line; ochre markings; copious pubescence inter- narrowing anteriorly and bordered pos- posed with longer setae; finely and densely teriorly; whitish pubescence with longer se- puncturated. Legs ochre with whitish setae; tae especially on the sides; coarsely and tibiae with numerous spines. Abdorrien densely puncturated. Elytra black or dark brown with ochre border of varying width; brown with ochre markings of varying size; stridulatory ridge marked, striae evident; copious pubescence interposed with longer post-metacoxal ridge complete; copious setae; with micropuncturation, and a coarser whitish pubescence with some longer se- irregular puncturation; striations evident. tae. Some males are hypermandibulated. Legs: femora and tibiae black, tarsi ochre; Similar species: A. sericans (Kiesenwetter). tibiae with numerous spines. Abdomen black or dark brown with ochre border; stridulatory Distribution ridge marked, striae evident; post-metacoxal Central Asia-Europe-Mediterranean region. ridge complete; copious whitish pubescence European distribution: southern Europe interposed with longer setae. Similar species: (WINKLER,1924; ZAITZEV,1924; CHARPENTIER,1965; A. pruinosus and A. curtus. MASCAGNI,1991); Transcaspia, Transcaucasia (ZAITZEV,1924); Corsica, Sardinia, Sicily Distribution (MASCAGNI,1985a); France, ex-Yugoslavia, Al- European distribution: central and north bania, Greece (MASCAGNI,1991); ltaly Europe (ZAITZEV,1924); France (BARTHE,1926); (MASCAGNI& CALAMANDREI,1992); Germainy, ltaly (PORTA,1929; MASCAGNI& CALAMANDREI, Switzerland, Austria, Hungary, Bulgaria, Tur- 1992); United Kingdom (CLARKE,1973); Eu- key (MASCAGNI,in press). rope (STRIEN,1980); Germany (MASCAGNI,1991). lberian records (fig. 19): Spain. Spain (KUWERT,1890; BARTHE,1926); Catalunya Studied material (fig. 19) (KIESENWETTER,1851, as minutus); Castellón: River Spain. Madrid: 1 ex., leg. Arias (MNCN); 1 ex., Mijares (MASCAGNI,1991); Sevilla (KIESENWE~ER,El Pardo, leg. Bolívar (MNCN); 1 ex., Aranjuez, 1865; FUENTE, 1929). coll. Pérez Arcas (MNCN).

Studied material (fig. 19) The species was not recorded from the Ibe- Spain. Southern Spain, 1 ex., leg. Schaufuss rian ~eninsula.According to BARTHE(1926), (MNCN); Castellón: 2 exs. 26 VI1 1954, River it is found in sandy substratum, both in Mijares, leg. A. Giordani Soika (MCVN), 1 ex. coastal areas and inland rivers. MASCAGNI same (CMS); Córdoba: 2 exs., coll. Pérez Arcas (1992, 1993) associated this species to the (MNCN); 5 exs., Posadas, coll. Bolivar (MNCN); sand~or rnudd~shores of running waters. Toledo: 8 exs., leg. Martínez (MNCN); 1 ex., coll. Pérez Arcas (MNCN). Augyles (Littorimus) maritimus (Guérin- According to BARTHE(1926) it is found in Méneville) Heterocerus rnaritimus Guérin-Méneville, Icon. the sandy shores of running waters, al- Regne anim. 1844: 69 Miscel.lania Zoologica 21.1 (1998) 79 Aguilera et al. nanus Gené, 1836 Ufer, leg. F. Hieke (MNB). Huesca: 2 exs. britannicus Kuwert, 1890 9 IV 1995, Barbastro, Cregenzán, River Ariño, burchanensis Schneider, 1896 leg. l. Ribera, G. Foster & P. Aguilera (CARB), basalis Mamitza, 1929 1 ex. same (CMS); 7 exs. 25 VI 1994, Fraga, melitensis Reitter, nomen nudum (Midsuf & Barranco de Valcuerna, leg. l. Ribera & P. Mascagni, 1997) Aguilera (CARB); 1 ex. 5 111 1994, 15 exs. 25 VI 1994, Pallaruelo de Monegros, Barranco Aedeagus as in figure 5. Total length 2.2- de Lafarda, leg. P. Aguilera & l. Ribera 3.4 mm. Head black, whitish pubescence in- (CARB), 1 ex. same (CMS). Murcia: 10 exs. terposed with longer setae especially on .the 19 111 1997, Lorca, Rambla de Torrealvilla, leg. sides; mandibles dentate with acute apex; l. Ribera & C. Hernando (CARB). Tarragona: clypeus with fine copious setae; antenriae 1 ex. IX 1933, Ampolla, leg. Museu (MZB); with 11 segments and 7-segmented club. 6 exs. 24 VI1 1994, Corbera d'Ebre, CN-420, Pronotum dark brown with reddish brown leg. l. Ribera & P. Aguilera (CARB), 1 ex. border and middle line; narrower anteriorly same (CMS); 2 exs. 24 VI 1995, Delta del Ebro, and bordered posteriorly; whitish pubescence La Marquesa, leg. P. Aguilera & l. Ribera with longer setae especially on the sides; (CARB), 1 ex. same (CMS); 1 ex. 27 V 1994, densely and finely puncturated. Elytra black Delta del Ebro, La Tancada, leg. P. Aguilera or dark brown with reddish markings of vary- (CARB); 4 exs., Valls, leg. Español (MZB). Va- ing size; copious pubescence interposed with lencia: l ex. 6 IV 1995, Quesa, leg. D. Lott & longer setae; micropuncturated and coarsely G. Foster (CARB), 1 ex. same (CFA). Zaragoza: and irregularly puncturated. Legs reddish 3 exs. 29 X 1996, Villafranca de Ebro, leg. P. brown; tibiae with numerous spines. Abdo- Aguilera (CARB). men black or dark brown sometimes with an ochre border; stridulatory ridge marked, striae Augyles maritimus is the most abundant evident; post-metacoxal ridge complete; co- and frequent species of the genus in the pious whitish pubescence interposed with lberian peninsula. It is mostly found in non- longer setae. Similar species: A. marmota vegetated shores of streams, or in the re- (Kiesenwetter). sidual pools left when temporary streams dry out. We have also found it in retro- Distribution dunal ponds and irrigation channels in the Europe - Mediterranean. Ebro Delta, in coexistence with Heterocerus European distribution: western and cen- flexuosus Stephens. It seems to tolerate a tral Europe (ZAITZEV,1924); France (BARTHE, wide range of salinity, being frequent in 1926); United Kingdom (CLARKE,1973); Sar- freshwater (e.g. in the head-waters of the dinia, Sicily (MASCAGNI,1985a); Mediterranean River Aguas, or in a residual pool in area (MASCAGNI,1991); ltaly (MASCAGNI& Corbera de Ebre), as well as in relatively CAMMANDREI,1992); Holland, Germany, Greece, mineralised waters (e.g. in Rambla de Malta (MASCAGNI,in press). Torrealvilla, Barranco de Lafarda, or the lberian records (fig. 20): Spain. Spain retro-dunal pool in La Tancada, in the Ebro (KUWERT,1890); Andalucía (BARTHE,1926); Va- Delta). It has not been found in the lencia: Albufera (MORODER,1924); Huesca: hypersaline Ramblas in southeastern Spain. Barbastro, Fraga, Pallaruelo de Monegros Despite its presence in inland localities in (RIBERAet al., 1996). France and the UK, it is considered a coastal species, associated to edges of rivers with Studied material (fig. 20) tidal influence, retro-dunal ponds, or salt Spain. Albacete: 1 ex. 16 111 1997, Paterna de water (BARTHE,1926; CLARKE,1973). The oc- Madera, Arroyo de la Fuenfría, leg. l. Ribera currence in the inland regions of the Ibe- & P. Aguilera (CARB). Almería: 8 exs. rian peninsula (habitually in semi-arid ar- 13 111 1997, 3 exs. 15 111 1997, Molinos de Río eas with mineralised waters) of species that Aguas, River Aguas, leg. l. Ribera (CARB); are considered to be exclusively coastal is a 10 exs. 6 IV 1996, Turre, River Aguas, leg. P. common pattern present in many groups, Aguilera (CARB). Cádiz: 1 ex. 7 V 1991, Alcalá as noted for aquatic Coieoptera by RIBERA de los Gazules (9 Km. SO), River Alberite- & AGUILERA(1995). Miscel.lania Zoologica 21.1 (1998)

Augyles (Littorimus) marmota (Kiesenwetter) According to BARTHE(1926) and MASCAGNI Heterocerus marmota Kiesenwetter, Stett. Ent. (1985b, 1992, 1993) it is associated to sandy Zeitg. XI, 1850: 224 or muddy shores. unicolor Dufour, 1851 funebris Schaufuss, 1862 maritimus Kuwert. 1890 Augyles (Littorimus) pruinosus (Kiesenwetter) Heterocerus pruinosus Kiesenwetter, Linnaea Ent. Aedeagus as in figure 4. Total length 2.2- V. 1851: 286 3.0 mm. Head black or dark brown, whitish pubescence with longer setae especially on Aedeagus as in figure 7. Total length 3.1- the sides; mandibles dentate, rather short, 4.0 mm. Head black, whitish pubescence in- with fairly acute apex; clypeus with fine terposed with longer setae; mandibles den- copious setae; antennae with 10 segments tate, rather short, with a fairly acute apex; and 6-segmented club. Pronotum dark clypeus with fine copious setae; antennae brown with reddish brown border and mid- with 11 segments and 7-segmented club. dle line; narrowing anteriorly and bordered Pronotum dark brown with reddish or red- posteriorly; whitish pubescence with longer dish brown border and middle line; narrow- setae especially on the sides; densely and ing anteriorly and bordered posteriorly; whit- finely puncturated. Elytra dark brown with ish pubescence with longer setae especially reddish markings of varying size; copious on the sides; densely and irregularly pubescence interposed with longer setae; puncturated. Elytra black with ochre mark- with micropuncturation, and a coarser and ings of varying size; copious pubescence in- irregular puncturation. Legs: femurs and terposed with longer setae; coarsely and ir- tibiae reddish brown, tarsi ochre, tibiae with regularly puncturated; striations evident. Legs: numerous spines. Abdomen dark brown, femurs and tibiae brown or reddish brown, sometimes with an ochre border; stridulatory tarsi ochre; tibiae with numerous spines. Ab- ridge marked, striae evident; post-metacoxal domen black or dark brown with ochre bor- ridge complete; copious whitish pubescence der; stridulatory ridge very marked, striae evi- interposed with longer setae. Similar spe- dent; post-metacoxal ridge complete; copi- cies: A. maritimus. ous whitish pubescence interposed with longer setae. Similar species: A. hispidulusand Distribution A. curtus. Europe-Mediterranean. European distribution: southern Europe Distribution (WINKLER,1924; ZAITZEV,1924); France (BARTHE, Europe. 1926); Sardinia, Sicily (MASCAGNI,1985a); European distribution: central Europe southern Europe, Greece (MASCAGNI,1991); (ZAITZEV,1924); France (BARTHE,1926); ltaly ltaly (MASCAGNI& CALAMANDREI, 1992); Roma- (PORTA,1929; MASCAGNI& CALAMANDREI, 1992); nia, Turkey (MASCAGNI,in press). central and southern Europe, western Sibe- lberian records (fig. 21): Girona (KIESENWEITER,ria (STRIEN,1980); Turkey (MASCAGNI,1991). 1850, 1851); western Pyrenees (FUENTE,1929). lberian records (fig. 21): eastern Pyrenees Spain. Spain (BARTHE,1926); central Spain (as (FUENTE,1929). Spain. Girona (FUENTE,1929); funebris, KUWERT,1890; FUENTE,1929); Málaga: Huesca: Fiscal (RIBERAet al., 1996). Cártama (ROSENHAUER,1856); Jaén, Sevilla (HEYDEN,1870; FUENTE,1929); Valencia: Albufera Studied material (fig. 21) (MORODER,1924), La Dehesa (TORRESSALA, 1962); Eastern Pyrenees, 1 ex., leg. J. Duval, coll. P. Alicante: El Llano (TORRESSALA, 1962). Arcas (MNCN). Spain. Girona: 1 ex. 21 V 1932, Pirineus, Sant Pau de Seguries, leg. Villarrubia Studied material (fig. 21) (MZB). Granada: 1 ex. (MNCN). Huesca: 2 exs. Spain. Andalucía: 1 ex., coll. Pérez Arcas 13 V 1995, Fiscal, River Ara, leg. P. Aguilera & (MNCN). Cuenca: 1 ex. VI1 1906, leg. Arias l. Ribera (CARB). (MNCN). Madrid: 3 exs., coll. Pérez Arcas (MNCN); 1 ex., Aranjuez (MNCN); 2 exs., El The locality in Huesca in the central pre- Pardo, leg. Bolívar (MNCN). Pyrenees was a permanent river with grave1 Aauilera et al. and boulders. The specimens were found by Studied material (fig. 22) washing the edges, among grave1 and vegeta- Spain. Almería: 1 ex. 5 IV 1996, River tion. In this station it coexisted with Heterocerus Almanzora, leg. P. Aguilera (CARB). Badajoz: holosericeus Rosenhauer. It seems to be a spe- 4 exs., leg. Uhagón (MNCN). Barcelona: 1 ex. cies preferentially found in mountains in the 11 VIII 1971, Balenya, leg. Villarrubia (MZB). lberian peninsula, so far only known from the Guadalajara: 9 exs. 25 IX 94, El Pobo de Pyrenees and probably Sierra Nevada (the stud- Dueñas, leg. P. Aguilera & l. Ribera (CARB), ied example in the MNCN had no details about 1 ex. same (CMS). Huesca: 2 exs. 25 VI 1994, the locality in which it was collected), although Pallaruelo de Monegros, Barranco de Lafarda, in ltaly it has been collected in a wider range leg. P. Aguilera & l. Ribera (CARB), 1 ex. of habitats (MASCAGNI,1992, 1993). same (CMS); 8 exs. 23 X 1994, Candasnos, leg. P. Aguilera (CARB), 1 ex. same (CMS). Jaén: 1 ex. 14 IV 1985, 1 km N Venta de la Augyles (Littorimus) senescens (Kiesenwetter) Nava, Puerto Zegrí, leg. G. N. Foster (CFA). Heterocerus senescens Kiesenwetter, Berl. Entom. Lleida: 1 ex. 23 VI 1990, Sanaüja, leg. l. Ribera Zeitschr. IX, 1865: 368, note (CARB). Madrid: 4 exs. (MNCN); 1 ex., Esco- punctatus Ch. Brisout de Barneville, 1866 rial (MNCN); 2 exs., Cercedilla (MNCN). Málaga: 2 exs. 16 IV 1985, 1 km S Río Grande Aedeagus as in figure 8. Total length 3.5- and Tolox, leg. G. N. Foster (CFA). León: 5.0 mm. Head black, whitish pubescence in- 1 ex., Ponferrada, leg. Paganetti (CSCR). terposed with very long setae; mandibles den- Sevilla: 1 ex. IV 1917 (MNCN); 6 exs., Lebrija, tate, rather short, with acute apex; clypeus leg. Snizek (CSCR), 1 ex. same (CMS); 4 exs. with fine copious setae; antennae with 11 seg- 11 V 1991, leg. Snizek (CSCR), 2 exs. same ments and 7-segmented club. Pronotum (CMS). black, narrowing anteriorly and bordered Portugal. Algarve: 1 ex. V 1910, Mon- posteriorly; whitish pubescence with copious chique, leg. Schatzmayr (CMS). Alto Alentejo: longer setae; coarsely and densely punc- 1 ex., Évora (MCM). turated. Elytra black with ochre markings of varying size; copious pubescence interposed It is habitually found in current waters, usu- with longer setae; micropuncturated and ally not mineralised (although it can occa- coarsely and irregularly puncturated; striations sionally occur in more salty waters, as in faint. Legs: femurs and tibiae black and ochre, Barranco de Lafarda in Huesca), of varied tarsi ochre; tibiae with numerous spines. Ab- flow and substratum. It has also been found domen black with ochre border; stridulatory in very recent ponds of rain water, in which ridge marked; post-metacoxal ridge complete; it was the only macroinvertebrate (e.g. El copious whitish pubescence interposed with Pobo de Dueñas, in Guadalajara), or coex- longer copious setae. isted with spp (Hydrophilidae) (e.g. Candasnos, Huesca). It seems thus to be Distribution a species with good colonising abilities. Western Europe. European distribution: Spain, Portugal (ZAITZEV,1924); France (BARTHE,1926); Sweden Augyles (Littorimus) sericans (Kiesenwetter) (GUILLEFORS,1982); Norway (MASCAGNI,in press). Heterocerus sericans Kiesenwetter, Germ. Zeitschr. IV, lberian records (fig. 22): Spain. Sevilla: 1843: 212, t. 3, f. 8 Castillejo (KIESENWETTER,1865); Spain (KUWERT, pusillus Stephens, 1835 1890; ZAITZEV,1924; BARTHE,1926); Santander: Reinosa (BRISOUTDE BARNEVILLE,1866 as H. Aedeagus as in figure 3. Total length 2.8- punctatus); Ciudad Real: Almuradiel, Sevilla: 3.6 mm. Head dark brown, whitish pubes- Castillejo (HEYDEN,1870); Badajoz, Ciudad Real, cence with long setae especially on the sides; Sevilla, Teruel (FUENTE,1929); Huesca: Can- mandibles dentate with acute apex; clypeus dasnos, Pallaruelo de Monegros (RIBERAet with fine copious setae; antennae with 11 seg- al., 1996). Portugal. Douro, Estremadura: ments and 7-segmented club. Pronotum dark Azambuja, Espinho (PAULINODE OLIVEIRA,1882, brown with dark orange border and fre- as punctatus; Zaitzev, 1924; FUENTE, 1929). quently a middle line; narrowing anteriorly Miscel.lania Zooloqica 21.1 (1998) 83

eo A. flavidus HA. hispidulus -150 km

Figs. 18-22. Distribution maps of the species of genus Augyles studied: m e Studied material; 0 Bibliographic data. (Records of wide geographical areas are not Aguilera et al.

and bordered posteriorly; whitish pubescence black and ochre, tarsi ochre; tibiae with nu- with some long setae especially on the sides; merous spines. Abdomen black with ochre finely and densely puncturated. Legs dark border of varying width; stridulatory ridge orange with whitish setae; tibiae with nu- marked, with very evident striae; post- merous spines. Abdomen dark brown, fre- metacoxal ridge incomplete; copious whitish quently with an ochre border; stridulatory pubescence with longer setae. Similar spe- ridge marked, striae evident; post-metacoxal cies: H. fenestratus Thunberg, H. fusculus ridge complete; copious whitish pubescence Kiesenwetter. with some longer setae. Similar species: A. flavidus. Distribution Western Mediterranean. Distribution European distribution: western Mediterra- Europe. nean (WINKLER,1924; ZAITZEV,1924; MASCAGNI, European distribution: central and east- 1991); France (BARTHE,1926); Sardinia, Sicily ern Europe, Transcaucasia (ZAITZEV,1924); (MASCAGNI,1985a); ltaly (MASCAGNI& &AMANDREI, France (BARTHE,1926); Yugoslavia, Euro- 1992); Germany (MASCAGNI,in press). turanian region (MASCAGNI,1991); ltaly lberian records (fig. 23): Girona (KIESENWETTER, (MASCAGNI& CALAMANDREI,1992); Great Brit- 1850,1851); eastern Pyrenees (FUENTE,1929). ain, Sweden, Holland (MASCAGNI,in press). Spain. Spain (KUWERT,1890, as mendax, as lberian records (fig. 22): Spain. Spain coxaepilus); Aragón (KUWERT,1890; FUENTE, (FUENTE,1929); Málaga: Cártama (ROSENHAUER,1929); Badajoz, Barcelona, Ciudad Real 1856, as minutus). (FUENTE,1929); Madrid: Escorial (KUWERT,1890 as mendax; FUENTE, 1929); Huesca: Barbastro The lberian examples of this species were (RIBERAet al., 1996); Girona: Capmany (RIBERA not studied. According to BARTHE(1926) and & AGUILERA,1996). Portugal. Portugal: MASCAGNI(1992) it is associated to sandy sub- "Amasia" (KUWERT,1890, as var. pictus; FUENTE, stratum. 1929), Alto Alentejo: Évora (MASCAGNI,1992).

Studied material (fig. 23) Tribu Heterocerini Spain. Cataluña: 1 ex., coll. Bolívar (MNCN). Galicia: 8 exs. (MNCN). Ávila: 2 exs. V 1934, Heterocerus Fabricius, 1792 Candeleda, River Tietar, leg. J. Baum (CSCR), 1 ex. same (CMS); 3 exs., Credos, Valle de Heterocerus aragonicus Kiesenwetter Iruelas, coll. Bolívar (MNCN); 1 ex., Navalperal Heterocerus aragonicus Kiesenwetter, Stett. Ent. (MNCN). Cádiz: 8 exs. 6 Vlll 1995, Jimena de Zeitg. XI, 1850: 223 la Frontera, River Hozgarganta, leg. P. pictus Mulsant & Rey, 1873 Aguilera (CARB), 1 ex. same (CMS). Castellón: coxaepilus Kuwert, 1890 2 exs. 6 IX 1897, 14 exs. 17 IX 1897, 8 exs. mendax Kuwert, 1890 VI 1898, 4 exs. 3 IV 1899, Fuente la Reina (MNCN). Córdoba: 4 exs., Posadas, coll. Aedeagus as in figure 15. Total length 3.5-5.0 Bolívar (MNCN). A Coruña: 2 exs., Malpica mm. Head black, whitish pubescence with (MNCN). Girona: 8 exs. 19 IX 1994, Bascara, longer setae especially on the sides; mandi- River Fluvia, leg. P. Aguilera & l. Ribera bles dentate with a fairly acute apex; clypeus (CARB), 3 exs. same (CMS); 81 exs. 9 VI1 1994, with fine copious setae; antennae with 11 seg- 67 exs. 10 IX 1994, Capmany, Riera de ments and 7-segmented club. Pronotum black Torrelles, leg. l. Ribera & P. Aguilera (CARB), with ochre border; narrowing anteriorly and 9 exs. same (CMS); 11 exs. 23 V 1991, La bordered posteriorly; whitish pubescence Janguera [La Jonquera?], leg. Snizek (CSCR), with longer setae especially on the sides; 1 ex. same (CMS). Huesca: 2 exs. 9 IV 1995, finely and densely puncturated. Elytra black Cregenzán, River Ariño, leg. l. Ribera, P. or dark brown with ochre markings of vary- Aguilera, G. Foster & D. Lott (CARB), 1 ex. ing size; copious pubescence interposed with same (CMS); 1 ex., Valle de Ordesa, leg. longer setae; micropunctures mixed with fine Bolívar (MNCN). Huelva: 8 exs. 9 V 1991, dense punctures. Legs: femurs and tibiae Santa Olalla del Cala (5 Km. N., 450 m.), leg. Misce1,lania Zoologica 21.1 (1998)

F. Hieke (MNB), 1 ex. sarne (CMS). Lleida: pusillus Waltl, 1839 1 ex. 26 V 1990, Oliana, River Segre, leg. l. tristis Mannerheim, 1853 Ribera (CARB). Madrid: 1 ex. (MCG); 2 exs. multimaculatus Motschulsky, 1854 (MCR); 2 exs. (MZF); 1 ex. VI 1995 (MNCN); fausti Reitter, 1879 1 ex., leg. G. Carrasco (MNCN); 1 ex. (MNCN); var. croaticus Kuwert, 1890 6 exs., Aranjuez (MNCN); 1 ex., Escorial, leg. siculus Kuwert, 1890 Cazurro (MNCN); 1 ex., Escorial (MNCN); var. corsicus Kuwert, 1890 10 exs., El Pardo, 610 m., coll. Sauffer fenestratus confinis Rey, 1890 (MNCN); 12 exs., El Pardo, coll. Sauffer fenestratus lanatus Rey, 1890 (MNCN); 3 exs., El Pardo, leg. Arias (MNCN); fenestratus obscurus Rey, 1890 1 ex., El Pardo (MNCN); 1 ex., Navacerrada, tonkinensis Grouvelle, 1896 leg. Uhagón (MNCN); 5 exs. V 1925, River latus Grouvelle, 1915 Alberche, leg. M. Escalera (MNCN); 3 exs. asiaticus Nomura, 1958 20 VI 1926, River Alberche (MNCN); 7 exs., completus Hatch, 1965 Sierra de Guadarrama, leg. J. Sauffer (MNCN). Oviedo: 2 exs. VI 1927, Villaviciosa, Aedeagus as in figure 6. Total length 2.5- leg. M. Escalera (MNCN), 4 exs. 10 VI 1927, 5.2 rnrn. Head black, whitish pubescence Villaviciosa (MNCN), 4 exs. 20 VI 1897, with longer setae especially on the sides; Villaviciosa (MNCN). Tarragona: 3 exs. rnandibles dentate, sornetirnes robust with 30 VI1 1931, Sant Vigens de Calders (MZB). acute apex; clypeus with fine copious setae; Valencia: 1 ex. 6 IV 1995, Millares, River antennae with 11 segrnents and 7-segrnent- Júcar, leg. D. Lott & G. Foster (CARB). ed club. Pronoturn black or dark brown Portugal. "Lusitania": 1 ex., coll. G. with ochre border of varying width; nar- Hauser, leg. Hurnrnler (MNB); Algarve: 2 exs. rowing anteriorly and bordered posteriorly; IV 1996, Ribeira da Boina, Pto. de Lagos, whitish pubescence with longer setae espe- leg. C. Hernando (CHB); Alto Alentejo: cially on the sides; finely and densely 5 exs., Évora, leg. A. Schatzrnayr (MCM), 2 exs. puncturated. Elytra black or dark brown same (MCT), 1 ex. sarne (CMS); Estrernadura: with ochre rnarkings varying greatly in size; 2 exs. IV 1996, Portalegre, River Caia, leg. C. copious pubescence interposed with longer Hernando (CHB). setae; rnicropunctures rnixed with fine dense punctures; striations faint. Legs: femurs and It is the rnost frequent and abundant spe- tibiae black and ochre, tarsi ochre; tibiae cies of the lberian Heterocerus. In the Ibe- with nurnerous spines. Abdomen black or rian peninsula it has been rnostly found in dark brown with ochre border of varying poorly rnineralised running waters with var- width; stridulatory ridge rnarked, striae evi- ied substraturn and vegetation, or in re- dent; post-rnetacoxal ridge incornplete; co- sidual pools in ternporary rivers or strearns pious whitish pubescence with longer se- (in agreernent with BARTHE,1926 and tae. Sorne of the males are hyperman- MASCAGNI,1985b. 1992, although the latter dibulated. Similar species: H. fusculus and also found the species in salt water). It can H. aragonicus. sometirnes forrn very populated colonies, e.g. in sandy shores or residual pools with- Distribution out vegetation in a ternporary strearn in Holarctic. Girona (Riera de Torrellas), where it was European distribution: the whole Palae- found with abundant larvae. The life cycle arctic region (WINKLER,1924; ZAITZEV,1924); and the irnrnature stages of the species were France (BARTHE,1926); England (CLARKE,1973); studied in detail by PIERRE(1946). Corsica, Sardinia, Sicily (MASCAGNI,1985a); Gerrnany, ex-Yugoslavia, Hungary, Greece, Turkey (MASCAGNI,1991); ltaly (MASCAGNI& (Thunberg) CALAMANDREI,1992). Dermestes fenestratus Thunberg, Nov. Acta Ups. lberian records (fig. 24): Spain. Girona: IV, 1784: 3 Rosas (CUN~,1885); Logroño (FUENTE,1929); laevigatus Panzer, 1794 Teruel: Calaceite (RIBERAet al., 1996). Portu- marshami Stephens, 1828 gal. Alto Alentejo: Évora (MASCAGNI,1992). 86 Aguilera et al.

Figs. 9-17. Male genitalia of the studied species of genus Heterocerus: 9. H. parallelus; 10. H. flexuosus; 11. H. fossor; 12. H. obsoletus; 13. H. holosericeus; 14. H. marginatus; Miscel.lania Zoologica 21.1 (1998) 87

E E E E E E m m m 2 2 2

15 16 17

Studied material (fig. 24) It is preferentially found in running waters, Spain. Alrnería: 7 exs. 6 IV 1996, Turre, River although it rnay occur in pools or in ponds Aguas, leg. P. Aguilera (CARB). Ávila: 5 exs., with abundant vegetation (e.g. in Aiguarnolls Navalperal (MNCN). Burgos: 15exs. 15 Vlll 1994, de I'Ernporda, or Aiguarnolls de Sils, in Villasandino, River Odra, leg. P. Aguilera Girona). In the lberian peninsula it has not (CARB), 1 ex. sarne (CMS). Cáceres: 1 ex. been found in highly rnineralized waters, al- 19 IV 1985, Abadía Largo, leg. G. N. Foster though according to BARTHE(1 926), CLARKE (CFA), Girona: 6 exs. 15 VI 1996, Aiguarnolls (1973) and MASCAGNI(1991) they constitute a de I'Ernporda, leg. P. Aguilera & C. Hernando typical habitat of this species. (CARB), 6 exs. sarne (CHB); 1 ex. 28 Vlll 1988, Aiguarnolls de Sils, leg. De Gregorio (MZB); 4 exs. V 1934, Ciurana, leg. Museu (MZB). HeterocerusflexuosusStephens Granada: 2 exs. 15 V 1989, Cubillas, leg. U. HeterocerusflexuosusStephens, III. Brit. 11, 1828: 101 Gollner (MNB), 1 ex. sarne (CMS); 3 exs. marginatusGyllenhal, 1808 23 V 1991; Girona: La Janguera [La Jon- femoralisKrynicky, 1832 quera?], leg. Snizek (CSCR), 1 ex. sarne (CMS). hamifer Gene, 1836 Madrid: 4 exs., Escorial (MNCN); 3 exs., Si- arenariusKiesenwetter, 1851 erra de Guadarrarna, leg. Sauffer (MNCN). maculosus Fairmaire, 1868 Oviedo: 1 ex. 20 VI 1897, Villaviciosa (MNCN). fausti Reitter, 1879 Tarragona: 30 exs. 1 V 1995, Caseres, River dentifasciatus Kuwert, 1890 Algars, leg. P. Aguilera (CARB), 1 ex. same damryiKuwert, 1890 (CMS). Teruel: 2 exs. 24 VI1 1994, Calaceite, heydeniKuwert, 1890 River Matarraña, leg. P. Aguilera & l. Ribera apfelbeckiKuwert, 1890 (CARB), 1 ex. sarne (CMS). corsicus Rey, 1890 Portugal. Algarve: 1 ex. IV 1996, Ribeira senegalensisFairmaire, 1894 da Boina, Puerto de Lagos, leg. C. Hernando (CHB). Alto Alentejo: 1 ex., Évora, leg. A. Aedeagus as in figure 10. Total length 3.5- Schatzrnayr (MCM), 4 exs. sarne (MCT). 5.6 rnrn. Head black, whitish pubescence with Aguilera et al. longer setae especially on the sides; mandi- P. Aguilera (CARB). Córdoba: 2 exs., leg. bles dentate, with acute apex; clypeus with Stempelmann (NMW), 1 ex. same (CMS). fine copious setae; male without clypeal horns; Málaga: 1 ex., Cártama, leg. Rosenhauer antennae with 11 segments and 7-segmented (MCVN). Murcia: 1 ex. 24 VI1 1994, Mar Menor, club. Pronotum black with brown or ochre S. Javier, leg. A. Giordani Soika (MCVN); 1 ex. border, narrowing anteriorly and feebly bor- 26 IV 1996, Salinas de San Pedro, leg. l. Ribera dered posteriorly; whitish pubescence with & A. Millán (CMM); 1 ex. 10 VI1 1992, longer setae especially on the sides; finely Cartagena, Marina de lo Poyo, leg. J.J. and densely puncturated. Elytra black with Sánchez Meca (CMM); 2 exs. 10 VI1 1992, ochre markings; copious pubescence inter- Cartagena, Lo Poyo, Los Nietos, leg. J. J. posed with longer setae; micropunctures Sánchez Meca (CMM). Sevilla: 1 ex. 11 V 1991, mixed with fine dense punctures. Legs ochre Lebrija (CSCR). Tarragona: 1 ex. VI 1934, with whitish setae; tibiae with numerous Alfacs, leg. Museu (MZB); 2 exs. 17 V 1994, spines and sometimes with fine black exter- Delta del Ebro, La Tancada, leg. l. Ribera, G. nal border. Abdomen black with ochre bor- Foster & D. Bilton (CARB), 2 ex. same (CFA), der of varying width; stridulatory ridge 1 ex. same (CMS); 3 exs. 24 VI 1995, Delta marked, with slight striae; post-metacoxal Ebro, La Marquesa, leg. P. Aguilera & l. Ribera ridge incomplete; copious whitish setae with (CARB); 8 exs. 24 VI 1995, Delta del Ebro, longer setae. Some of the males are hyper- L'Alfacada, leg. P. Aguilera & l. Ribera (CARB); mandibulated. Similar species: H. fossor 1 ex. VI 1934, Sant Carles de la Rapita, leg. Kiesenwetter and H. parallelus Gebler. Museu (MZB). Baleares: 3 exs., Mallorca, Rabassa, leg. Breit (MCM); 2 exs., Mallorca, Distribution leg. W. Pepp, ex coll. Winkler (MCG), 3 exs. Palaearctic. same (MCVN), 1 ex. same (CMS). European distribution: central Europe (ZAITZEV,1924); France (BARTHE,1926); United The species is habitually found in salt marshes Kingdom, lreland (CLARKE,1973); Corsica, Sar- with sand or muddy substratum (BARTHE,1926; dinia, Sicily, ex-Yugoslavia; Cyclad lslands CLARKE,1973; MASCAGNI,1985b, 1992). In the (Greece) (MASCAGNI,1985a); Greece, Ural lberian peninsula it is also frequently found mountains, Caucasus (MASCAGNI,1991); ltaly in coastal lagoons with different degrees of (MASCAGNI& CALAMANDREI, 1992); Norway, Swe- salinity, sometimes directly connected to the den, Finland, Denmark, Holland, Germany, sea (e.g. I'Alfacada, Tarragona, in a retro- Czech Republic, Switzerland, Austria, Hun- dunal pond with infiltrated sea water, with gary, Romania, Bulgaria, Albania, Cyprus, a fauna formed mostly by marine macro- Turkey (MASCAGNI,in press). invertebrates). The species is usually found lberian records (fig. 25): eastern Pyrenees in sandy substratum, although it also occurs (FUENTE,1929). Spain. Spain (KUWERT,1890, as in grave1 or coarse sand (e.g. in Torreblanca, femoralis); southern Spain (KUWERT,1890, as Castellón). It has sometimes been found in dentifasciatus; FUENTE,1929); Alicante: Calpe coexistence with Augyles maritimus. (MORODER.1924; Torres Sala, 1962); Valencia: Godelleta (MORODER,1924); Murcia: Nlar Menor, Málaga: Cártama (MASCAGNI,1991). Heterocerus fossor Kiesenwetter Balearic Islands: Ibiza. Salinas (ESPANOL,1943). Heterocerus fossor Kiesenwetter, Germ. Zeitschr. Mallorca (MASCAGNI,1985a; MASCAGNI,1991). IV, 1843: 204, t. 3, f. 3 bifasciatus Kuster, 1849 Studied material (fig. 25) parallelus Dufour, 1852 Spain. Alicante: 1 ex. 23 Vlll 1994, Santa Pola, rectus Waterhouse, 1859 Salinas, leg. P. Aguilera & l. Ribera (CARB), fossor ab. cincticollis Rey, 1890 1 ex. same (CMS); 1 ex. 29 11 1992, El Hondo, leg. J.J. Sanchez Meca (CMM). Barcelona: Aedeagus as in figure 11. Total length 4.5- 2 exs. VI 1931, Llobregat (MZB). Castellón: 6.2 mm. Head black, whitish pubescence with 10 exs. 27 Vlll 1981, Oropesa de Mar, leg. M. longer setae especially on the sides; mandi- Rizzotti (MCVR), 2 exs. same (CMS); 5 exs. bles dentate, very robust with acute apex; 12 1 1997, Torreblanca, PIa de Cabanes, leg. male with clypeal horns; clypeus with fine Miscel.lania Zoologica 21.1 (1 998) copious setae; antennae with 11 segments Aedeagus as in figure 17. Total length 2.4- and 7-segmented club. Pronotum black with 3.8 mm. Head black, whitish pubescence with ochre border, narrowing anteriorly and fee- longer setae especially on the sides; mandi- bly bordered posteriorly; whitish pubescence bles dentate with acute apex; clypeus with with longer setae especially on the sides; fine copious setae; antennae with 11 seg- finely and densely puncturated. Elytra black ments and 7-segmented club. Pronotum with ochre markings of varying size; copious black or dark brown with dark orange bor- pubescence interposed with longer setae; der; narrowing anteriorly and bordered finely and densely puncturated; striations posteriorly; whitish pubescence with longer evident. Legs ochre with whitish setae; tibiae setae especially on the sides; finely and with numerous spines and with black exter- densely puncturated. Elytra black or dark nal border. Abdomen black with ochre bor- brown with orange markings of varying size; der of varying widths; stridulatory ridge copious pubescence interposed with longer marked, striae evident; post-metacoxal ridge setae; micropunctures mixed with fine dense incomplete; copious whitish pubescence with punctures; striations very faint. Legs dark longer setae. Some of the males are brown with brown tarsi; tibiae with numer- hypermandibulated. Similar species: H. ous spines. Abdomen black or dark brown flexuosus and H. parallelus. with orange border of varying width; stridulatory ridge marked, striae evident; Distribution post-metacoxal ridge incomplete; copious Euro-Siberian. whitish pubescence with longer setae. Simi- European distribution: central and south- lar species: H. fenestratus. ern Europe, south-western Siberia (ZAITZEV, 1924); France (BARTHE,1926); Sardinia (PORTA, Distribution 1929); England, lreland (CLARKE,1973); EU- Euro-Siberian. rope, southern Siberia (STRIEN,1980); Germany, European distribution: Europe, western Si- Turkey (MASCAGNI,1991); ltaly (MASCAGNI& beria (WINKLER,1924; ZAITZEV,1924); France CA~MANDREI,1992); Sweden, Finland, Holland, (BARTHE,1926); ltaly (PORTA,1929; MASCAGNI& Belgium, Czech Republic, Austria, Hungary, CALAMANDREI,1992); England (CLARKE,1973); Romania, Bulgaria, Greece (MASCAGNI,in press). Germany, ex-Yugoslavia (MASCAGNI,1991); lberian records: Spain (FUENTE,1929). Sweden, Finland, Holland, Belgium, Poland, Balearic lslands (fig. 26): Mallorca, La Czech Republic, Switzerland, Austria, Roma- Puebla, Pollensa (TENENBAUM,1915). nia (MASCAGNI,in press). lberian records (fig. 26): Portugal. Beira Studied material (fig. 26) Litoral: Estarreja (PAULINODE OLIVEIRA,1882; Portugal. Alto Alentejo: 2 exs., Évora, leg. A. FUENTE,1929). Schatzmayr (MCM). It is a European species, whose presence in These are the first Portuguese records of the lberian peninsula is in need of confirma- the species. In Britain it is always associated tion. The only lberian record was published with salt water areas of clay and mud (CLARKE, in a general catalogue, not in a specialized 1973), although in other geographical ar- work (PAULINODE OLIVEIRA,1882). According eas it can also be found in the shores of to CLARKE(1973). in England it is exclusively freshwater rivers (BARTHE,1926; MASCAGNI, found on the south coast, at the base of 1985b, 1992). sea-cliffs in muddy ground. In France and ltaly it frequently occurs in sandy river banks, although it can also be found in more min- Heterocerus fusculus Kiesenwetter eralized waters (BARTHE,1926; MASCAGNI,1992). Heterocerus fusculus Kiesenwetter, Gerrn. Zeitschr. IV, 1843: 220, t. 3, f. 1 1 oblongulus Kuwert, 1890 Heterocerus holosericeus Rosenhauer pulchellus Kuwert, 1890 Heterocerus holosericeus Rosenhauer, Thier. Andal. sirnilis Kuwert, 1890 1856: 114 fusculus var. opacus Rey, 1890 panorrnitanus Kuwert, 1890 Aguilera et al. ragusae Kuwert, 1890 Studied material (fig. 27) ragusae var. lineatus Kuwert, 1890 Spain. Southern Spain: 1 ex. (MNCN). holosericeus var. pustulatus Schilsky, 1890 Andalucía: 2 exs. (MNCN). Almeria: 8 exs. pustulatus Schilsky, 1892 15 Vlll 1994, Mojacar, River Aguas, leg. P. villiger Fiori, 1906 Aguilera (CARB), 1 ex. same (CMS); 2 exs. fluviatilis Fiori, 1906 13 111 1997, Molinos de Río Aguas, River subfossor Fiori, 1906 Aguas, leg. l. Ribera (CARB). Barcelona: 1 ex. villosus Fiori. 1908 9 IV 1932, Balenya, leg. Villarrubia (MZB). Cádiz: 3 exs., Chiclana, coll. Pérez Arcas Aedeagus as in figure 13. Total length 4.0- (MNCN). Granada: 1 ex. 18 VI1 1945, Puerto 5.0 mm. Head black, whitish pubescence with Camacho, leg. E. Zaico (MNCN). Huesca: 2 exs. numerous and longer setae; mandibles den- 13 V 1995, Fiscal, River Ara, leg. P. Aguilera & tate, rather short, with a fairly round apex; l. Ribera (CARB). Málaga: 1 ex., Cártama, leg. clypeus with fine copious setae; antennae with Rosenhauer (MCVN). Tarragona: 2 exs., Valls, 11 segments and 7-segmented club. Pronotum leg. Español (MZB). black, with reddish border; narrowing anteriorly and bordered posteriorly; whitish In coincidence with BARTHE(1926) and pubescence with longer setae especially on MASCAGNI(1985b, 1992), the species has been the sides; fine dense punctures. Elytra black found in the shores of running waters, both with reddish markings of varying size; copious in the head-waters (e.g. River Aguas en pubescence interposed with longer setae; Molinos, Almería, or River Ara in Fiscal, micropunctures mixed with fine dense punc- Huesca) and areas next to the coast (e.g. tures; striations evident. Legs reddish brown, River Aguas in Mojacar). It has been found tarsi ochre; tibiae with numerous spines. Ab- in coexistence with Augyles pruinosus and domen black with reddish border of varying A. maritimus. It is also habitually found in width; stridulatory ridge marked, striae evi- flood detritus (MASCAGNI,198513). dent; post-metacoxal ridge incomplete; copi- ous whitish pubescence with longer setae.

Distribution Heterocerus marginatus (Fabricius) Europe -Mediterranean. Apate marginatus Fabricius, Mant. Ins. 1, 1787: 33 European distribution: western Mediter- sulcatus Kuwert, 1890 ranean region (WINKLER,1924; CHARPENTIER,scutellaris Guillebeau, 1897 1965); Spain, North Africa (ZAITZEV,1924); Sic- ily (ZAITZEV,1924; MASCAGNI,1985a); France Aedeagus as in figure 14. Total length 3.5- (BARTHE,1926); Europe, western Siberia (STRIEN, 4.2 mm. Head black, whitish pubescence with 1980); ltaly (ZAITZEV,1924; PORTA,1929; longer setae especially on the sides; mandibles MASCAGNI& CALAMANDREI,1992); Turkey dentate, rather short with acute apex; clypeus (MASCAGNI,1991). with fine copious setae; antennae with 11 seg- lberian records (fig. 27): Spain. Spain ments and 7-segmentedclub. Pronotum black, (KUWERT,1890; ZAITZEV,1924; BARTHE,1926); sometimes with a reddish border; narrowing Málaga: Cártama, Guadalhorce (ROSENHAUER,anteriorly and bordered posteriorly; whitish 1856; FUENTE, 1929; MASCAGNI,1991); Córdoba, pubescence with longer setae especially on Sevilla: Guadajoz (KIESENWEITER,1865; HEYDEN, the sides; micropunctures mixed with fine dense 1870; FUENTE, 1929); Valencia: River Júcar, punctures. Elytra black with reddish markings Carcaixent, La Parrilla (MORODER,1924), La of varying size; copious pubescence interposed Dehesa, Carcaixent (TORRESSALA, 1962); with longer setae; micropunctures mixed with Alicante: El Llano (TORRESSALA, 1962); Huesca: fine dense punctures; striations evident. Legs Fiscal (RIBERAet al., 1996). Portugal. reddish brown, tarsi ochre; tibiae with numer- "Felgueira", Beira Litoral: Coimbra, Douro, ous spines. Abdomen black or dark brown Douro Litoral: River Vizela (PAULINODE Oliveira, with ochre border of varying widths; stridu- 1882; FUENTE, 1929). latory ridge marked, striae evident; post- Balearic Islands: Mallorca, Col d'en Rabasa metacoxal ridge incomplete; copious whitish (TENENBAUM,1915). pubescence with longer setae. Miscel.lania Zoologica 21.1 (1998)

Disitribution Distribution Palaearctic. Euro-Siberian. European distribution: Europe, south- European distribution: central and south- western Siberia (ZAITZEV,1924); France ern Europe, south-western Siberia (ZAITZEV, (BARTHE,1926); England (CLARKE,1973); Sar- 1924); France (BARTHE,1926); England (CLARKE, dinia, Sicily (MASCAGNI,1985a); Euro- 1973); Corsica, Sicily (MASCAGNI,1985a); from turanian region, Austria (MASCAGNI,1991); Siberia to the Turkestan, Italy, Austria ltaly (MASCAGNI& CALAMANDREI, 1992); Nor- (MASCAGNI,1991); Sweden, Finland, Denmark, way, Sweden, Denmark, Holland, Belgium, Holland, Belgium, Germany, Hungary, Greece, Germany, Poland, Czech Republic, Ucrania, Turkey (MASCAGNI,in press). Switzerland, Romania, ex-Yugoslavia, Bul- lberian records: Spain (KUWERT,1890; FUENTE, garia, Greece (MASCAGNI,in press). 1929) (without detailed localities). lberian records (fig. 28): Spain. Southern lberian examples of this species were not Spain (KUWERT,1890; as sulcatus; BARTHE,1926); studied. In Britain and France it is found in Valencia: Albufera (MORODER,1924); Barce- salt water (BARTHE,1926; CLARKE,1973). lona: Hospitalet (Cu~i& MARTORELL,1876), Ciudad Real (FUENTE,1929). lberian examples of this species were not Heterocerus parallelus Gebler found. It seems to occur habitually in water Heterocerusparallelus Gebler, Bemerk. Ins. Sib. in bodies associated to freshwater rivers Ledeb. Reise 11, 1830, App. 111: 101 (BARTHE,1926; CLARKE,1973). although in salinus Kiesenwetter, 1843 MASCAGNI(1992) it is also recorded from salt rnaxillosus Motschulsky, 1854 water. According to DAJOZ(1965), it flies cornutus Motschulsky, 1854 readily to UV light. hauseri Kuwert, 1893

Aedeagus as in figure 9. Total length 4.8- 7.6 mm. Head black, whitish pubescence with Heterocerus obsoletus Curtis longer setae especially on the sides; mandi- Heterocerusobsoletus Curtis, Brit. Entorn. V. 1828: 224 bles dentate, very robust, with a fairly acute rnarginatus Marsham, 1802 apex, dorsal projection and no clypeal horns obsoletus var. quadrirnaculatus Hochhuth, 1873 in males; smaller, with a fairly acute apex, and no dorsal projections in females; clypeus with Aedeagus as in figure 12. Total length 4.2- fine copious setae; antennae with 11 segments 5.8 mm. Head black, whitish pubescence with and 7-segmented club. Pronotum black with numerous and longer setae; mandibles den- ochre border of varying width, narrowing tate and robust with acute apex; clypeus with anteriorly and feebly bordered posteriorly; fine copious setae; antennae with 11 seg- whitish pubescence with longer setae espe- ments and 7-segmented club. Pronotum cially on the sides; finely and densely black, with reddish border; narrowing puncturated. Elytra black with ochre mark- anteriorly and bordered posteriorly; whitish ings of varying size; copious pubescence in- pubescence with very longer setae espe- terposed with longer setae; finely and cially on the sides; micropunctures mixed densely puncturated. Legs ochre with whit- with fine dense punctures. Elytra black or ish setae; tibiae with numerous spines. Ab- dark brown with reddish markings of vary- domen black with faintly ochre border; ing size; copious pubescence interposed stridulatory ridge marked, without evident with very long setae; micropunctures mixed striae; post-metacoxal ridge incomplete; with with fine dense punctures; striations evi- copious whitish pubescence with longer se- dent. Legs brown; tibiae with numerous tae. Some males are hypermandibulated. Simi- spines. Abdomen brown with reddish bor- lar species: H. fossor and H. flexuosus. der of varying width; stridulatory ridge marked, without evident striae; post- Distribution metacoxal ridge incomplete; copious whit- Euro-Siberian. ish pubescence with longer setae. European distribution: southern Europe, 92 Aguilera et al.

H. aragonicus -150 km

bibliográficos. (No se incluyen citas de ámbito geográfico general. Las citas provinciales de las que no se conoce la localidad precisa se sitúan sobre la capital de la provincia.) Miscel.lania Zoologica 21.1 (1998) 93

South-western Siberia (ZAITZEV,1924); France Micilus (M. murinus Kiesenwetter, 1843) is also (BARTHE,1926); Austria, ex-USSR (MASCAGNI, present (MASCAGNI,1992). France, with 10 1991); ltaly (MASCAGNI& CALAMANDREI, 1992); Heterocerus, 11 Augyles and Micilus murinus Germany, Poland, Hungary, Bulgaria, Greece (ZAIREV,1924; BARTHE,1926; MASCAGN~,in press) (MASCAGNI,in press). apparently has a richer fauna, with al1 lberian lberian records (fig. 29): Spain. Southern species plus A. aureolus Schiodte, 1866, A. Spain (KUWERT,1890; FUENTE,1929); Barcelona: humilis Rey, 1890 (with a dubious taxonomic Casa Antúnez, Hospitalet, Bordeta (CUN~& validity), A. intermedius (Kiesenwetter, 1843), MARTORELL,1876; Cu~i,1888; FUENTE,1929). one subspecies, H. fusculus ssp. pulchellus Kiesenwetter, 1843, and M. murinus. Studied material (fig. 29) In Morocco 13 species have so far been Spain. Spain: 1 ex., leg. G. Carrasco (MNCN). recorded, six Augyles (one, A. humilis, not Cádiz: 3 exs., Algeciras (MNB), 1 ex. same (CMS). present in the lberian peninsula, although-the record is considered to be dubious by KOCHER, According to BARTHE(1926) it is associated to 1956) and seven Heterocerus (two of them not salt waters. found in Iberia, H. biskrensis Fairmaire, 1879 and H. dayremi Peyerimhoff, 1921) (KOCHER, 1956). Two additional species are known from Discussion Algeria, H. pallidivestis Guillebeau, 1893 and H. humilis Guillebeau, 1896 (= H. guillebeaui Of the 17 species of Heteroceridae present in Zaitzev, 1908) (~AIREV,1924). the lberian peninsula and the Balearic lslands As regards the fauna of the Balearic 1s- (eight Augyles and nine Heterocerus), 16 oc- lands, and in comparison with other Medi- cur in Spain (the only missing species is H. terranean Islands, four species are known fusculus, whose presence in lberia is in need from Corsica, eight from Sardinia (two pend- of confirmation), seven in Portugal (two ing confirmation), and 11 from Sicily (one of Augyles and seven Heterocerus), and only them dubious) (MASCAGNI,1985a) (note the three in the Balearic lslands (three in Mallorca synonymisation of A. gravídus, H. hamifer and one in Ibiza) (table 1). In comparison and H. siculus by MASCAGNI,in press). with nearby faunas, Italy, despite being bet- The number of species and the composi- ter studied, has the same number of species tion of the heterocerid fauna in Spain, despite of Augyles and Heterocerus, with the only the scarcity of data, is very similar to that of difference being the absence of A. senescens, nearby geographical areas. Both in Portugal and the presence of A. crinitus (Kiesenwetter, and the Balearic lslands it is most likely that 1850). In ltaly the only European species of additional species would be found if properly 94 Aguilera et al.

Species

...- ...... -.. .- ... -. ...- .. -- ... . -. ...-. .. .. - .-. .-. . -. -...... -..... -. . A. marmota (Kiesenwetter, 1850) + + ...... -..... -...... - ...... -. .-. .- -. .- -..... -. -. . -.. - ......

H. fossor Kiesenwetter, 1843 + + + .. -- ...- -. .- -...... - ...... U...... -. ... - .. - ......

searched, specially taking into account the the holotype and one recent record from a wide distribution of most European species. nearby locality RIBERA^^ al., 1977). Its general The absence of endemic species of this family distribution is therefore still uncertain. How- in western Europe stands out, in coincidence ever, in family Elmidae there are some endemic with what happens in family Dryopidae (OLMI, species with restricted distributions, in particu- 1976). There are only two known endemic lar in the genus Oulimnius (OLMI, 1976; lberian species of this family, Dryops hispanus BERTHÉLEMY,1979). Knowledge of the fourth Olmi, 1973 (of a more than dubious taxo- lberian family of Dryopoidea, Limnichidae, is nomic validity, Eugenio Rico, personal com- almost non-existent, being reduced to the few munication, 1996). and D. championi Dodero, records included in FUENTE(1929). All the known 1918. The latter species is only known from species of western European heterocerids have Miscel.lania Zoologica 21.1 (1998) 95 a wide distribution.This could be partly due humilis, known from southern France, and to their high dispersa1 abilities (although in Micilus murinus, widely distributed in Eu- other groups there are flying species with very rope, and present in Grenoble and Lyon restricted distributions, see e.g. NOONAN,1990). (BARTHE, 1926). Some of the north African Possible species to be searched in the Ibe- species are also likely present in the south- rian peninsula are those with a more north- east of Spain, dueto the close biogeographi- ern European distribution, in particular A. cal links between these two areas. 96 Aguilera et al.

a middle line reddish brown, densely and finely puncturated. Elytra dark brown with

finely punctura;ed. Elytra without striaiions 7 . . .. -...... - . -- . . -. .- . . . - .. -. .- ...... - - . . - .. . -.. . - .. .. .

and a middle line dark orange. Elytra dark brown with ochre markinas, finelv and denselv

middle line reddish brown. Elytra black or dark brown with reddish markings, coarsely and ir-

tal length 2.2-3.4 mm. Aedeagus as in figure 5 rnaritirnus -. . ------. - - .- . - . .. -. .. - - . . .. . - .. -- -. . -. . . - -. - - . .- . . 8. Elytra brown or reddish brown. Leqs ochre. To- tál length 3.0-3.5 mm. Aedeagus as in figure 1 curtus Elytra black or dark brown. Legs brown or

9. Pronotum dark brown, with the border and a middle line reddish or reddish brown. Miscel.lania Zoologica 21.1 (1998) 97 Aauilera et al.

Acknowledgements References

We thank Giorgio Alberti (MCT), Miguel BARTHE,E., 1926. Tableaux analytiques des Ángel Alonso Zarazaga (MNCN), Luca Coléopteres de la faune franco-rhénane, Bartolozzi (MZF), Oleguer Escola (MZB), Garth famille XLVII, Heteroceridae. Misc. Ent., 29: Foster (CFA), Carles Hernando (Barcelona), 1-32. Fritz Hieke (MNB), Isabel Izquierdo (MNCN), BERTH~LEMY,C., 1979. Elmidae de la region Manfred Jach (MNW), Jose Luis Lencina Palearctique occidentale: systematique et (Jumilla), Carlo Leonardi (MCM), Dereck Lott repatition (Coleoptera Dryopoidae). Annls (Leicester), Andrés Millán (Murcia), Giuseppe Limnol., 15: 1-102. Osella (Dip. Sc. Ambient. Univ. I'Aquila), BRISOUTDE BARNEVILLE,CH., 1866. Coléopteres Roberto Poggi (MCG), Enrico Ratti (MCVN), nouveaux trouvés en Espagne pendant Juanjo Sánchez Meca (Cartagena), Stanislav I'excursion de la societé en 1865. Ann. Skalicky (Czech Republic), Manfred Uhlig Soc. Ent. Fr., 6: 355-426. (MNB) and Vincenzo Vomero (MCR) for al- CHARPENTIER,R., 1965. A monograph of the lowing us to study their material. We also family Heteroceridae (Col.) of the Ethio- thank Garth Foster and Franck Bameul for pian region. South African life, their comments on the manuscript. 11: 215-243. CLARKE,R. Q. S., 1973. Handbooks for the identification of British . Coleoptera Resumen Heteroceridae. Royal Entomological Soci- ety, London. La familia Heteroceridae MacLeay, 1825 CROWSON,R. A., 1960. The phylogeny of (Coleoptera, Dryopoidea) en la península Coleoptera. Ann. Rev. Entom., 5: 11 1-134. Ibérica e islas Baleares Cu~i,M., 1885. Especies de insectos que he descubierto recientemente en Cataluña. En este trabajo se revisa toda la información Actas Soc. esp. Hist. nat., 14: 70-73. disponible sobre la familia Heteroceridae - 1888. Insectos observados en los alrede- en la península Ibérica e islas Baleares, dores de Barcelona. An. Soc. esp. Hist. basándose en datos publicados y en el nat., 17: 133-191. estudio de material de colecciones o el Cu~i,M. & MARTORELL,M., 1876. Catálogo me- recolectado por los autores. Se figuran los tódico y razonado de los coleópteros ob- edeagos de las 17 especies estudiadas (figs. servados en Cataluña. Imprenta Gorchs, 1-17). ocho Augyles y nueve Heterocerus, y Barcelona. se incluye una breve descripción de todas Dnioz, R., 1965. Faune terrestre et d'eau douce ellas. Se cita Augyles hispidulus Kiesenwetter des Pyrénées-Orientales. Fascicule 9. Cata- por primera vez en la península Ibérica, en logue des Coléopteres de la foret de la España, y Heterocerus fossor Kiesenwetter Massane. Masson & Cie., Paris. por primera vez en Portugal. Se incluye ESPANOL,F., 1943. Contribución al conocimien- una clave de los géneros europeos y las to de los Coleópteros de Ibiza y Formen- especies iberobaleares de la familia. Se tera y un nuevo Heliophilus ibérico (Col. incluyen algunas breves notas ecológicas, Tenebrionidae). Las Ciencias, 8: 93-108. junto con los mapas de distribución en la FUENTE, J. M. DE LA, 1929. Catálogo sistemáti- Península y las Baleares (figs. 18-29). co-geográfico de los Coleópteros obser- Aunque el número de especies hasta el vados en la Península Ibérica, Pirineos pro- momento citadas de España (ocho Augyles piamente dichos y Baleares (continuación). y ocho Heterocerus) es similar al de otras Bol. Soc. ent. Esp., 12: 84-99. zonas geográficas próximas, la fauna de GUILLERFORS,C., 1982. Heterocerus senescens, Portugal, con solo siete especies, y la de las ny skalbaggsart fort Sverige. Ent. Tidskr., islas Baleares, con solo tres, se pueden 103: 71-72. considerar todavía como escasamente HEYDEN,L. VON, 1870. Entomologi~cheReise conocidas. Se discute la posible presencia nach dem südlichen Spanien der Sierra en España de especies de distribución Guadarrama und Sierra Morena, Portugal europea o norteafricana. und der Cantabrischen Gebirgen. Berlin. Miscel.lania Zooloqica 21.1 (1998)

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