The Coleopterists Bulletin, 61(1):95–110. 2007.

DIVERSITY OF WATER (COLEOPTERA:GYRINIDAE,HALIPLIDAE, NOTERIDAE,HYGROBIIDAE,, AND HYDROPHILIDAE) IN GALICIA, NORTHWEST SPAIN:ESTIMATING THE COMPLETENESS OF THE REGIONAL INVENTORY

JAIME GONZA´ LEZ Departamento de Bioloxı´a , Faculdade de Bioloxı´a Universidade de Santiago de Compostela 15706 Santiago de Compostela, SPAIN [email protected]

ANDRE´ S BASELGA Departamento de Biodiversidad y Biologı´a Evolutiva Museo Nacional de Ciencias Naturales c/ Jose´ Gutie´rrez Abascal 2 28006 Madrid, SPAIN [email protected]

AND

FRANCISCO NOVOA Departamento de Bioloxı´a Animal, Faculdade de Bioloxı´a Universidade de Santiago de Compostela 15706 Santiago de Compostela, SPAIN [email protected]

Abstract Diversity of Galician water beetles (Coleoptera: Gyrinidae, Haliplidae, Noteridae, Hygrobiidae, Dytiscidae and Hydrophiliade) was analyzed using a database created by collecting species records from several data sources. Most data came from a long-term sampling program carried out by one of the authors across the whole region but further local studies provided additional data. To assess the inventory completeness as well as to estimate the species richness, both an asymptotic model and some non-parametric estimators were used. Database records were utilized as a sampling-effort surrogate. Total richness estimations predicted by these different methods range between 117 and 131 species. Therefore, it seems that between 86 and 97% of the fauna belonging to the studied families was recorded, so that the inventory has reached an acceptable level of completeness (113 species). A list of the Galician species of Gyrinidae, Haliplidae, Noteridae, Hygrobiidae, Dytiscidae and Hydrophilidae is given.

Water beetles are an important part of the populations of water bodies and wetlands. They are used as indicators of ecological diversity and habitat conservation (Foster 1987; Eyre and Foster 1989; Foster et al. 1990; Ribera and Foster 1993; Sa´nchez-Ferna´ndez et al. 2004) as they meet most of the criteria generally accepted in the selection of indicator taxa (Pearson 1994) and are especially useful in certain habitats as peat bogs, coastal and saline lagoons, wood and wetland ponds, etc (Ribera and Foster 1993). Because of the importance of water bodies and wetlands in the planning of conservation efforts, the assessment of the completeness of water beetle

95 96 THE COLEOPTERISTS BULLETIN 61(1), 2007 inventories can be considered an essential task. There is a great deal of papers discussing how to use several methods as a way to estimate the species richness for a wide range of taxonomic groups. The use of randomized sample accumulation curves was first developed for standardized samplings (Sobero´n and Llorente 1993; Colwell and Coddington 1994; Carlton and Robison 1998; Moreno and Halffter 2000; Summerville et al. 2001; Noguera et al. 2002). However, these methods were also applied to different types of data coming from non- standardized samplings, like museum collections or faunistic databases (Sobero´n et al. 2000; Hortal et al. 2001; Martı´n-Piera and Lobo 2003; Petersen et al. 2003; Rosenzweig et al. 2003; Hortal et al. 2004; Meier and Dikow 2004; Baselga and Novoa 2006). Another way to determine the degree of completion of the species inventories in a region is to plot the growth throughout time of the cumulative species number as a function of the year of description or first record (Medellı´n and Sobero´n 1999; Cabrero-San˜udo and Lobo 2003; Baselga and Novoa 2006). The knowledge of the water beetle diversity in the Iberian Peninsula has been increasing distinctly throughout the last 25 years, thanks to the contributions of a number of researchers who produced monographs on several areas. Ribera (2000) estimates the number of Iberian species of the families studied here falling between 263 and 280. The first records known for Galicia are from the second half of the 19th century (Heyden 1870). During the first half of the 20th century only a few specimens were captured and it is not until 1980 that several studies, considering either the whole region or some smaller areas, were undertaken. An extensive research carried out by Gonza´lez (1993) in the whole region produced an inventory of 94 species. Further studies referring to smaller areas have raised this number to the present 113 species and have improved significantly the knowledge of its distribution. The aim of this paper is to assess the advance in the knowledge of the diversity of the Galician fauna of water beetles by determining the degree of completion of the species inventory.

Material and Methods All the families of aquatic fall under the scope of this study as well as aquatic species of the family Hydrophilidae. The remaining families of water beetles were not included due to the lack of data. Most data used herein come from collecting done by Gonza´lez between 1984 and 1991 in a large survey of the autonomous community of Galicia, in the NW of Spain (Fig. 1) (Gonza´lez and Novoa 1988; Gonza´lez et al. 1989; Gonza´lez 1993; Gonza´lez and Novoa 1995). This resulted in 209 sampling sites and 94 species. The remaining records come from further studies carried out in smaller areas throughout the last 20 years. Garrido and Re´gil (1989) studied the valley of the Limia river, in the south, and produced a list of 46 species; Garrido (1990), Garrido and Re´gil (1994) provided some records from the western spurs of the Cantabrian range; De Paz and Otero (1993) undertook a complete study of the Landro river, in the north, and produced an inventory of 32 species; Novoa et al. (2004) studied the Fragas do Eume Natural Park and found 30 species; Garrido and Sa´inz-Cantero (2004) found 51 species in their work at Serra do Barbanza and Gonza´lez et al. (2005) inventoried 47 species from Serra do Xistral. All these areas have been marked in Fig. 2, which shows the spatial distribution of the sampling sites. In the former study most of the sampling sites were visited only once while in the later ones every site was visited several times throughout the sampling period (Fig. 3). THE COLEOPTERISTS BULLETIN 61(1), 2007 97

Fig. 1. Location of the studied area in the Northwest of the Iberian Peninsula.

In all 750 samples, from 305 sites, were obtained. A total of 25,297 specimens were captured in those samples which have been deposited in the Gonza´lez and De Paz collections at the University of Santiago de Compostela and in the Garrido collection at the University of Vigo. Some specimens, captured by Luis Iglesias throughout the first half of the 20th century, and identified by one of the authors (Gonza´lez 1993), are deposited in the Luis Iglesias Museum at the University of Santiago de Compostela. A few records by other researchers (Chapman and Champion 1907; Iglesias 1928; Bertrand 1956; Lagar 1961; Hansen 1982; Berge Henegouwen 1986; Fery and Heindrich 1988; Fresneda et al. 1990; Fery 1991; Angus et al. 1992; Balke and Fery 1993; Fery and Brancucci 1997; Fery 1999) were also considered. To assess the completeness of the inventory both an asymptotic model and some non-parametric estimators (ICE, Chao 2, jackknife of first and second order) were used (Colwell and Coddington 1994). The accumulation curve and the non-parametric estimators were generated with EstimateS 7.0 software (Colwell 2004), randomizing the sample order 100 times. Database records were used as a sampling-effort surrogate (Sobero´n et al. 2000; Hortal et al. 2001; Martı´n-Piera and Lobo 2003). Our database comprised 3,312 records for 25,314 specimens of 113 species. Each record is composed of the following fields: species name, site, date, number of specimens and collector. Any difference in any database field value gives rise to 98 THE COLEOPTERISTS BULLETIN 61(1), 2007

Fig. 2. Spatial distribution of sampling sites. The squared pattern corresponds with the UTM grid. Each 100 km2 square with one or more sampling sites is shown in grey. Areas with intense sampling (see text for details) are marked with diamonds. a new database record, so that increments of the number of records provide correlative increments of the sampling effort (Martı´n-Piera and Lobo 2003). Thereafter, the asymptotic Clench function was fitted to the smoothed curve (Sobero´n and Llorente 1993; Hortal et al. 2004):

SeðÞ~ ae=ðÞ1 z be where S(e) is the number of species found per sampling effort unit e; a and b, the parameters of the function. The latter were adjusted to the data of each curve by means of a Simplex and Quasi Newton method (StatSoft 2001). The predicted asymptote is calculated as a/b. The second estimation to determine the degree of completeness of the Galician water beetles inventory was produced by fitting the Clench function to the cumulative number of recorded species from the year of the first records in the THE COLEOPTERISTS BULLETIN 61(1), 2007 99

Fig. 3. Sampling frequencies. Most sites were sampled only one or two times. Sites sampled more than two times belong to the areas marked with diamonds in (Fig. 2). region (Heyden 1870) to the present, following (Cabrero-San˜udo and Lobo 2003) but considering the first Galician record instead of the year of description. This historic curve was generated taking into account the year of publication of the papers or the year of collection if known. The final section (1980–2004) of this curve seems to get an asymptotic shape due to an increase in the sampling effort and thus only this period was selected to adjust the Clench function.

Results The accumulation curve generated out of the database records (Fig. 4) reaches a species richness (S obs 5 113) very close to the asymptotic value predicted by the Clench function (116.6) (R2 of the adjusted function 5 0.99) and not far from the non-parametric estimators ICE (120.6), Chao 2 (122.4), Jackknife 1 (125.0) and Jackknife 2 (131.0). The inventory of aquatic Adephaga and Hydrophilidae seems, therefore, to have reached a remarkable level of completeness, since the referred sampling campaigns (combined dataset) found out between 86 and 97% of the Galician species of water beetles. It is expected that further studies will only find a few more species still not recorded. When the historic growth of the specific richness of Galician water beetlesis analyzed (Fig. 5a) it can be noticed that over 80% of the species have been collected in a short period of time, from 1980 to the present, when an important increase of the sampling effort was made. The curve becomes asymptotic during this period (Fig. 5b) so that it is possible to adjust the Clench function (R2 5 0.95) and to obtain a new estimation of the total species richness which should be expected in the area. The asymptote corresponding to this historic curve is 130.8 and, therefore, the estimated degree of completeness is 86%. To verify if the completeness was homogeneous in both groups, aquatic Adephaga and Hydrophilidae, two different accumulation curves were generated (Fig. 6). The Clench function does not adjust to the Hydrophilidae curve. This 100 THE COLEOPTERISTS BULLETIN 61(1), 2007

Fig. 4. Species accumulation curve generated from the database with the fitted Clench function and the non-parametric estimators ICE, Chao2 and 1st and 2nd order jackknife. suggests an underestimation of the species richness, so that the calculated completeness of 99% seems rather unrealistic and thus useless if we consider the non-parametric estimators. Each estimation for the two groups is shown in Table 1 and these results suggest a more complete inventory for aquatic Adephaga than for Hydrophilidae.

Discussion We consider the Galician inventory of the water beetle families studied herein to be reasonably complete, especially in the case of aquatic Adephaga. The lack of completion observed in the case of Hydrophilidae may be due to the characteristics of the sampling program which was more specific for aquatic Adephaga. Besides, data coming from (De Paz and Otero 1993; Garrido and Re´gil 1989) do not include this later family.

Table 1. Estimations for Hydradephaga (S obs 5 85) and Hydrophilidae (S obs 5 28).

Group Hydradephaga Hydrophilidae Estimator S S obs/S S S obs/S Clench Asymptote 88.4 96% 28.4 99% ICE 87.8 97% 33.6 83% Chao 2 88.0 97% 31.6 89% Jack 1 91.0 93% 33.0 85% Jack 2 93.0 91% 36.0 78% THE COLEOPTERISTS BULLETIN 61(1), 2007 101

Fig. 5. a) Historic curve (1870–2004) (dots) together with the number of species collected each year (bars). b) Fitted Clench function considering the final period (1980– 2004). 102 THE COLEOPTERISTS BULLETIN 61(1), 2007

Fig. 6. a) Species accumulation curve using only Hydradephaga records. b) Species accumulation curve using only Hydrophilidae records. THE COLEOPTERISTS BULLETIN 61(1), 2007 103

The Galician inventory of aquatic Adephaga and Hydrophilidae (113 species) represents 43% of the Iberian diversity of these families. This value is similar to that found for leaf beetles (Baselga and Novoa 2006) and slightly lower than that attained for ground beetles (A. Campos, pers. comm.), probably because this family includes a considerable number of Galician endemics. The species richness (113 taxa) observed in Galicia (29,574 km2) is comparable to those from other Iberian regions as Segura River basin (14,432 km2) where 102 species were found or Arago´n (47,720 km2) where 149 species were recorded. The historic curve shows the poor knowledge of Galician water beetles before 1980. The extensive study carried out by Gonza´lez (1993) involved a substantial contribution, increasing the known richness to 94 species, with further local studies increasing the total species number by 13%, as well as improving our knowledge about species distributions. It is remarkable that all the estimations obtained in this paper fluctuate between a narrow range. Both asymptotic models based either on the accumulation of database records or on the increment of the historic richness, and the non-parametric estimators assess richness values between 117 and 131 species. Even if we evaluate the completeness of aquatic Adephaga and Hydrophilidae inventories separately, the addition of its respective estimations is almost identical to the values yielded by the combined analysis. The coincidence of the estimations computed using different methods indicates that these values can be considered reliable. One possible bias that could influence our results could be the accumulation of a great number of repetitive records coming from local studies. The effect of such a reiteration in oversampled sites could yield underestimations for the Galician richness value. To assess this possible bias, we have removed the most reiterative records, coming from (De Paz and Otero 1993; Garrido and Re´gil 1989). The database is then reduced to 1908 records for 110 species, but the estimations are almost the same, both using the Clench asymptote (116.0) and the non-parametric estimators (120.6–133.0). Therefore, it seems that the reiteration effect of some sites is not biasing our results. Another possible biasing source are the sampling methods. Though estimates diverge in narrow ranges, as mentioned above, the only exception are the Hydrophilidae with estimates more heterogeneous. This is possibly indicative of a sampling bias produced by the use of collecting methods more appropriate for aquatic Adephaga. Finally, it was compared the effectiveness in estimating richness values of an extensive sampling performed by a single collector against the complete dataset. The field work conducted by Gonza´lez between 1984 and 1991 has the advantage of the homogeneity in samplings, making possible the use of samples (different day/locality) as sampling effort surrogate. Using this reduced data set (7,264 specimens, 1,354 database records, 323 samples, 94 species) the estimates are not realistic since they fluctuate between 99 and 106, all of them lower than the current number of recorded species (113). Exactly the same values are obtained both using database records or samples as sampling effort surrogate (Clench 5 102, ICE 5 99, Chao 2 5 100, Jack 1 5 105, Jack 2 5 106). As a concluding remark, the importance of faunistic studies carried out since 1980 should be emphasized. This work has increased the inventory of aquatic Adephaga and Hydrophilidae from 26 species collected before 1980 (though only 7 were published) to the 113 species known at present, allowing current good knowledge about Galician water beetle diversity and thus making the present analysis on the completeness of the inventory possible, as well as the estimations of actual species richness. 104 THE COLEOPTERISTS BULLETIN 61(1), 2007

Field work and faunistic papers are a basic background for any biodiversity research, a fact that was previously exemplified for Galician Chrysomelidae (Baselga and Novoa 2006). This paper showed that predictions on leaf beetles species richness result in strongly biased underestimations when using data coming from studies lacking extensive sampling effort. No research on invertebrate biodiversity should be conducted without the essential workof extensive field sampling and the publication of faunistic inventories, especially for hyper-diverse groups as Coleoptera.

Acknowledgments This study was supported by Xunta de Galicia with projects PGDIT01P- XI2004PR and PGDIT01MAM20001PR. Andre´s Baselga position at Museo Nacional de Ciencias Naturales (Madrid) is supported by Juan de la Cierva Program (MEC).

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Appendix A. A checklist of the Galician water beetles of the families Gyrinidae, Haliplidae, Noteridae, Hygrobiidae, Dytiscidae and Hydrophilidae. The year of the first capture is given when known. Otherwise the publication year appears.

Species First record Reference Remarks Gyrinidae Gyrinus (Gyrinus) caspius 1987 Gonza´lez and Novoa, Me´ne´trie´s, 1832 1995 Gyrinus (Gyrinus) substriatus 1914 Gonza´lez and Novoa, Stephens, 1829 1995 Gyrinus (Gyrinus) urinator 1921 Fuente, 1921 Illiger, 1807 Orectochilus villosus (O. F. 1984 Gonza´lez and Novoa, Mu¨ller, 1776) 1995 Haliplidae Peltodytes caesus (Duftschmid, 1995 Garrido and Sa´inz- 1805) Cantero, 2004 Peltodytes rotundatus 1981 Gonza´lez and Novoa, (Aube´, 1836) 1995 Haliplus (Neohaliplus) lineatocollis 1980 Gonza´lez and Novoa, (Marsham, 1802) 1995 Haliplus (Haliplus) heydeni 1981 Gonza´lez and Novoa, Whencke, 1875 1995 Haliplus (Liaphlus) guttatus 1985 Garrido and Re´gil, 1989 Aube´, 1836 Haliplus (Liaphlus) rubidus 1986 Gonza´lez and Novoa, Perris, 1857 1995 Noteridae Noterus laevis Sturm, 1834 1981 Gonza´lez and Novoa, 1995 Hygrobiidae Hygrobia hermanni (Fabricius, 1980 Gonza´lez and Novoa, 1775) 1988 Dytiscidae Liopterus atriceps Sharp, 1882 1984 Garrido and Re´gil, 1989 Liopterus haemorrhoidalis 1981 Gonza´lez and Novoa, (Fabricius, 1787) 1988 Laccophilus hyalinus 1980 Gonza´lez and Novoa, (Aube´, 1837) 1988 Laccophilus minutus (Linnaeus, 1981 Gonza´lez and Novoa, 1758) 1988 Hyphydrus aubei Ganglbauer, 1981 Gonza´lez and Novoa, 1891 1988 Hydrovatus clypealis Sharp, 1981 Gonza´lez and Novoa, 1876 1988 Yola bicarinata (Latreille, 1804) 1985 Garrido and Re´gil, 1989 Bidessus coxalis Sharp, 1882 1986 Gonza´lez and Novoa, 1995 Bidessus goudotii (Laporte, 1835) 1981 Gonza´lez and Novoa, 1988 Bidessus minutissimus (Germar, 1985 Garrido and Re´gil, 1824) 1989 Hydroglyphus geminus (Fabricius, 1981 Gonza´lez and Recorded as H. 1792) Novoa, 1988 pusillus (Fabr.) Hygrotus (Coelambus) lagari 1986 Gonza´lez and Novoa, Recorded as Coelambus (Fery, 1992) 1988 parallelogrammus (Ahrens) Hygrotus (Coelambus) 1984 Gonza´lez and Novoa, marklini (Gyllenhal, 1813) 1988 108 THE COLEOPTERISTS BULLETIN 61(1), 2007

Appendix A. Continued. Species First record Reference Remarks Hygrotus (Hygrotus) inaequalis 1981 Gonza´lez and Novoa, (Fabricius, 1777) 1988 Hydroporus decipiens Sharp, 1877 1984 Gonza´lez and Novoa, Recorded as H. analis 1988 (Aube´) Hydroporus brancoi brancoi 1979 Fery, 1999 Rocchi, 1981 Hydroporus brancuccii Fery, 1987 1989 Gonza´lez, 1993 Hydroporus discretus Fairmaire & 1984 Gonza´lez and Novoa, Brisout, 1859 1988 Hydroporus foveolatus Heer, 1839 1986 Garrido, 1994 Hydroporus gyllenhalii Schio¨dte, 1980 Gonza´lez and Novoa, 1841 1995 Hydroporus necopinatus 1985 Gonza´lez and Novoa, Recorded as H. cantabricus necopinatus Fery, 1999 1988 Sharp Hydroporus nevadensis Sharp, 1882 1990 Gonza´lez and Novoa, Recorded as H. longulus 1995 Mulsant Hydroporus nigrita (Fabricius, 1981 Gonza´lez and Novoa, 1792) 1988 Hydroporus normandi normandi 1984 Garrido and Re´gil, 1989 Re´gimbart, 1903 Hydroporus obsoletus Aube´, 1838 1984 Garrido and Re´gil, 1989 Hydroporus paganettianus Scholz, 1999 Novoa et al. , 2003 1923 Hydroporus planus (Fabricius, 1986 Garrido, 1994 1781) Hydroporus pubescens (Gyllenhal, 1981 Eiroa et al., 1988 1808) Hydroporus sabaudus (Fauvel, 1995 Garrido and Sa´inz- Recorded as H. nivalis Heer 1865) Cantero (2004) Hydroporus tessellatus Drapiez, 1956 Bertrand, 1956 1819 Hydroporus vagepictus Fairmaire & 1921 Gonza´lez and Novoa, Laboulbe`ne, 1854 1988 Hydroporus vespertinus Fery & 1981 Gonza´lez and Novoa, Galician records of H. Hendrich, 1988 1995 erytrocephalus, now excluded form the Iberian fauna, are included here Graptodytes aequalis Zimmermann, 1995 Garrido and Sa´inz- 1918 Cantero, 2004 Graptodytes flavipes Olivier, 1981 Gonza´lez and Novoa, Recorded as G. concinnus 1795 1988 Steph. Graptodytes ignotus (Mulsant & 1984 Garrido and Re´gil, Rey, 1861) 1989 Graptodytes varius (Aube´, 1838) 1984 Gonza´lez and Novoa, 1988 Scarodytes halensis (Fabricius, 1984 Garrido and Re´gil, 1787) 1989 Stictonectes epipleuricus 1981 Gonza´lez and Novoa, (Seidlitz, 1787) 1988 Stictonectes lepidus (Olivier, 1785) 1979 Gonza´lez and Novoa, 1988 Deronectes angusi Fery & 1986 Gonza´lez and Novoa, Recorded as D. latus Brancucci 1990 1988 (Stephens) Deronectes bicostatus (Schaum, 1986 Gonza´lez and Novoa, 1874) 1988 Deronectes costipennis gignouxi 1984 Gonza´lez and Novoa, Recorded as D. hispanicus. Fery & Brancucci, 1989 1988 Later, D. costipennis costipennis Brancucci was also found in Galicia (Gonza´lez et al., 1989) THE COLEOPTERISTS BULLETIN 61(1), 2007 109

Appendix A. Continued. Species First record Reference Remarks Deronectes ferrugineus Fery & 1986 Garrido, 1990 Brancucci, 1987 Deronectes opatrinus (Germar, 1984 Garrido and Re´gil, 1989 1824) Stictotarsus bertrandi (Legros, 1978 Gonza´lez and Novoa, 1956) 1988 Stictotarsus duodecimpustulatus 1980 Gonza´lez and Novoa, (Fabricius, 1792) 1988 Stictotarsus griseostriatus (De 1987 Garrido, 1990 Geer, 1774) Nebrioporus (Nebrioporus) 1985 Gonza´lez and Novoa, carinatus (Aube´, 1838) 1988 Nebrioporus (Nebrioporus) elegans 1985 Gonza´lez and Novoa, (Panzer, 1794) 1988 Nebrioporus (Zimmermannius) 1984 Garrido and Re´gil, ceresyi (Aube´, 1838) 1989 Oreodytes sanmarkii alienus (Sharp, 1984 Gonza´lez and Novoa, Recorded as Oreodytes 1872) 1988 rivalis Gyllenhal Agabus (Agabus) labiatus (Brahm, 1985 Garrido and Re´gil, 1791) 1989 Agabus (Gaurodytes) biguttatus 1973 Gonza´lez and Novoa, Recorded as A. nitidus (Olivier, 1795) 1988 (Fabr.), now considered a synonym Agabus (Gaurodytes) bipustulatus 1914 Gonza´lez and Novoa, (Linnaeus, 1767) 1988 Agabus (Gaurodytes) brunneus 1984 Gonza´lez and Novoa, (Fabricius, 1798) 1988 Agabus (Gaurodytes) conspersus 1986 Gonza´lez and Novoa, (Marsham, 1802) 1988 Agabus (Gaurodytes) didymus 1937 Gonza´lez and Novoa, (Olivier, 1795) 1988 Agabus (Gaurodytes) guttatus 1907 Chapman and guttatus (Paykull, 1798) Champion, 1907 Agabus (Gaurodytes) heydeni 1963 Fresneda et al, 1990 Wehncke, 1872 Agabus (Gaurodytes) nebulosus 1985 Garrido and Re´gil, (Forster, 1771) 1989 Agabus (Gaurodytes) paludosus 1984 Garrido and Re´gil, (Fabricius, 1801) 1989 Ilybius chalconatus (Panzer, 1796) 1986 Garrido, 1990 Ilybius dettneri (Fery, 1986) 1963 Fresneda et al., 1990 Ilybius meridionalis (Aube´, 1837) 1924 Gonza´lez, 1993 Galician records of I. fuliginosus (Fabr.), a species confined to the Pyrenees, are included here Ilybius montanus (Stephens, 1828) 1961 Lagar Mascaro´, 1961 Recorded as Agabus melanocornis Zimmermann Rhantus (Rhantus) hispanicus 1981 Gonza´lez and Novoa, (Sharp, 1882) 1988 Rhantus (Rhantus) suturalis 1980 Gonza´lez and Novoa, Recorded as R. pulverosus (McLeay, 1825) 1988 (Steph.) Colymbetes fuscus (Linnaeus,1798) 1980 Gonza´lez and Novoa, 1988 Acilius (Acilius) sulcatus 1985 Garrido and Re´gil, (Linnaeus, 1758) 1989 marginalis marginalis 1924 Gonza´lez and Novoa, Linnaeus, 1758 1988 Dytiscus pisanus Laporte, 1835 1926 Gonza´lez and Novoa, 1988 110 THE COLEOPTERISTS BULLETIN 61(1), 2007

Appendix A. Continued. Species First record Reference Remarks Dytiscus semisulcatus O. F. Mu¨ller, 1924 Gonza´lez and Novoa, 1776 1988 Cybister (Scaphinectes) 1928 Gonza´lez and Novoa, lateralimarginalis (De Geer, 1988 1774) Hydrophilidae Berosus (Berosus) affinis Brulle´, 1980 Gonza´lez (1993) 1835 Berosus (Berosus) hispanicus 1995 Garrido and Sa´inz- Ku¨ster, 1847 Cantero (2004) Berosus (Berosus) signaticollis 1980 Gonza´lez (1993) (Charpentier, 1825) Chaetarthria seminulum seminulum 1986 Gonza´lez (1993) (Herbst, 1797) Paracymus scutellaris (Rosenhauer, 1921 Gonza´lez (1993) 1856) Anacaena (Anacaena) bipustulata 1980 Gonza´lez (1993) (Marsham, 1802) Anacaena (Anacaena) globulus 1981 Gonza´lez (1993) (Paykull, 1798) Anacaena (Anacaena) lutescens 1987 Gonza´lez (1993) (Stephens, 1829) Anacaena (Anacaena) limbata 1928 Iglesias (1928) (Fabricius, 1792) Laccobius (Hydroxenus) femoralis 1991 Gonza´lez (1993) Rey, 1885 Laccobius (Dimorpholaccobius) 1984 Gonza´lez (1993) atratus Rottenberg, 1874 Laccobius (Dimorpholaccobius) 1981 Gonza´lez (1993) obscuratus Rottenberg, 1874 Laccobius (Dimorpholaccobius) 1986 Gonza´lez (1993) sinuatus Motschulsky, 1849 Laccobius (Dimorpholaccobius) 1905 Gentili (1988) striatulus (Fabricius, 1801) Laccobius (Dimorpholaccobius) 1921 Gonza´lez (1993) ytenensis Sharp, 1910 Helochares (Helochares) lividus 1870 Heyden (1870) (Forster, 1771) Helochares (Helochares) punctatus 1981 Gonza´lez (1993) Sharp, 1869 Enochrus (Lumetus) fuscipennis (C. 1982 Gonza´lez (1993) G. Thomson, 1884) Enochrus (Lumetus) halophilus 1923 Gonza´lez (1993) (Bedel, 1878) Enochrus (Methydrus) nigritus 1981 Gonza´lez (1993) (Sharp, 1872) Cymbiodyta marginella (Fabricius, 1995 Sa´inz-Cantero and 1792) Garrido (1996) Hydrobius convexus Brulle´, 1835 1995 Garrido and Sa´inz- Cantero (2004) Hydrobius fuscipes (Linnaeus, 1981 Gonza´lez (1993) 1758) Limnoxenus niger (Zschach, 1788) 1928 Iglesias (1928) Hydrochara flavipes (Steven, 1808) 1996 Garrido and Sa´inz- Cantero (2004) Hydrophilus (Hydrophilus) 1986 Gonza´lez (1993) pistaceus (Laporte, 1840) Coelostoma (Coelostoma) 1987 Gonza´lez (1993) hispanicum (Ku¨ster, 1848) Coelostoma (Coelostoma) 1868 Heyden (1870) orbiculare (Fabricius, 1775)