Harpalus (Baeticoharpalus) Lopezi, New Subgenus and New Species of Harpalus from the Iberian Peninsula (Coleoptera, Carabidae: Harpalini)

Harpalus (Baeticoharpalus) lopezi, new subgenus and new species of Harpalus from the Iberian Peninsula (Coleoptera, Carabidae: Harpalini)

José Serrano & José Luís Lencina

Serrano, J. & Lencina, J. L. 2008: Harpalus (Baeticoharpalus) lopezi,newsub- genus and new species of Harpalus from the Iberian Peninsula (Coleoptera, Ca- rabidae: Harpalini). – Entomol. Fennica 19: 193–198. In this paper we describe the new subgenus Baeticoharpalus Serrano & Lencina and a new species Harpalus (Baeticoharpalus) lopezi Serrano & Lencina, based on a unique character combination that includes the pubescence of temples, frons, pronotum and elytra, the lack of mentum tooth, the shape of mouth parts and the male genitalia. The new taxon was collected in Sierra Morena (Pico de Almadén, central Betic Mountains, south Spain) and is geographically isolated from related taxa. Distribution data suggest that this is a relict species. The taxo- nomic position of the new taxon is discussed in relation to the other Iberian subgenera of Harpalus and the related genera Pangus, Pseudoophonus, Ophonus, etc. A key to Iberian Harpalus and related taxa is included; it illustrates the abun- dance of homoplasic characters within these taxa and the difficulties to develop a ‘natural’ key for identifying them. J. Serrano & J. L. Lencina, Department of Zoology and Physical Anthropology, University of Murcia, Facultad de Veterinaria, Campus de Espinardo, 30071 Murcia, Spain; E-mails: [email protected] & [email protected] Received 15 February 2007, accepted 29 October 2007

1. Introduction Noonan (1976) lumped most taxa related to Harpalus in a single genus, whereas Kataev et al. The genus Harpalus Latreille, 1802 is one of (2003) rised Ophonus Dejean, 1821, Pangus De- largest of the family Carabidae in species number. jean, 1821 and Typsiharpalus Tshitschérine, 1901 According to Noonan (1976) it includes 32 sub- (among other taxa) to the genus rank. Other taxa genera and about 480 species mostly in the Pale- such as Cryptophonus Brandmayr & Zetto- arctic, Nearctic and Neotropical Regions; there Brandmayr, 1982 and Pseudoophonus Motschul- are 60 Ethiopic species and a few ones in the Ori- sky, 1844 are currently recognized as subgenera ental Region. In the recent World list of Lorenz of Harpalus, except for authors following the (2005), the genus includes fewer subgenera (10) Jeannel school. and species (367), because some of the subgenera The separation of Harpalus from related gen- admitted by Noonan were ranked as genera. The era and the division of this genus into subgenera genus shows a remarkable variability in external has not yet attained a general agreement. There is characters which explains the different criteria no World key for all these taxa, nor a Palearctic put forward by authors about its limits. Thus, key of the subgenera of Harpalus. Important 194 Serrano & Lencina • ENTOMOL. FENNICA Vol. 19 characters to differentiate Ophonus and related 2. Baeticoharpalus subg. n. taxa from Harpalus and its allied groups were discussed by Brandmayr et al. (1980), including Type species: Harpalus (Baeticoharpalus) lopezi the type of clypeous teeth, the setae on the stipes Serrano & Lencina, sp. n., by monotypy and the supernumerary teeth of mandibles in the Diagnosis. Black, shiny, without dorsal larvae, the dorsal vestiture, the pubescence of microsculpture. Size around 9 mm. Wingless. paraglossae and the aedeagus in the imagoes. Antennae infuscate except for the pale red first These characters led Brandmayr et al. (1980) to antennomere, and reaching the elytral basis. Palpi distinguish an ophonoid lineage made up by all pale red. Short pubescence present in temple the subgenera of Ophonus and the Harpalus (some isolated setae are also found in the frons), tenebrosus group (this last was ranked as the new sides and margins of pronotum, and in all elytral subgenus Cryptophonus by Brandmayr and intervals. Pronotum transverse, slightly sinuate Zetto-Brandmayr in 1982), from the harpaloid before the posterior angles, that are obtuse but lineage that includes Harpalus, Pseudoophonus, well marked. Scutellar stria present of average Pardileus, Acardystus, and Harpalophonus length. Humeri evident with a small tooth. Elytra among other taxa. The analysis of these charac- slightly sinuate before apex. Femora and tibiae al- ters showed that apomorphies defining the most black, tarsi paler, pro- and mesotarsi of ophonoid lineage are only found in the larvae, be- males dilated. Tarsi glabrous above except for the cause key characters of the adult seem to be nota- normal apical setae. Paraglossae pubescent, first bly homoplasic as they have been developed in- labial palpomere without diagonal carina, ligula dependently in separate lineages of Harpalus and with two apical setae, mentum without tooth. related taxa (Brandmayr et al. 1980). The molec- Submentum, prosternum, proepisternae and pro- ular data reported by Martínez-Navarro et al. pleura with short pubescence, mesosternum gla- (2005) showed the isolation of Ophonus from the brous and almost impunctate. Metaepisternae Harpalus-related taxa, the affinity between Pseu- longer than wider (L/W = 1.2–1.4), moderately doophonus and Harpalus and the equivocal posi- punctate and pubescent. Metasternum glabrous tion of Cryptophonus. except for 1–2 isolated setae. Last ventral ster- When attempting to identify five individuals nites pubescent at each side of median line. Me- of a hairy Harpalus species collected in Sierra dian lobe of aedeagus average for the genus, with Magina (central Betic Mountains, south Spain), it a straight apex moderately dilated and discrete was clear that they corresponded to a new spe- apical disk; the sclerites (two teeth, one field of cies. Further, the combination of external charac- microtrichia and another of macrotrichia) are also ters did not match those of currently accepted average for the genus. subgenera existing in the western Palearctic re- Specific epithet. The new subgenus is named gion, and thus the new species possibly repre- according to the location of the type locality in the sented an unknown lineage deserving the subge- Betic sierras. nus rank. These preliminary findings were quite Remarks. The new subgenus shows a number surprising because the genus Harpalus is well of affinities to other Harpalus, as are the general known in the Iberian Peninsula, and the Betic form and size, the details of mouth parts (pubes- Mountains have been reasonably sampled in the cence of paraglossae, labial palpi), and the type of last decades. This paper describes the new spe- aedeagus. However, the pubescence of temples cies, discusses the erection of a new subgenus of and frons combined with the lack of mentum Harpalus and formulates a number of hypotheses tooth are not found in subgenera of Harpalus. about the new species to be verified in future The new subgenus is geographically isolated as studies, including larval and molecular analyses. no other species of the west Palearctic Region A key to Iberian Harpalus and related taxa is in- shows these features. However, its relationships cluded to reflect the problems derived from the should be further investigated by analyzing usage of homoplasic characters in the lineage of Harpalus taxa from Asia, where the genus is Harpalus sensu lato, and to help in evaluating the much diversified (see Kataev 1987, 1991, 2001 systematic position of the new species. and further references therein). In particular, it ENTOMOL. FENNICA Vol. 19 • Baeticoharpalus, a new carabid subgenus 195 becomes clear that dorsal vestiture, including the temples and the frons, is a homoplasic character not suitable for phylogenetic analyses in Harpa- lus and related taxa. Likewise, the diagonal carina of the first labial palpomere is also homoplasic as it is found in both the ophonoid (all members) and the harpaloid (Pseudoophonus) lineages. This compels to elaborate key to adults based on com- binations of multiple characters instead of using apomorphies (see the key below). It is hypothesized that larvae of Baetico- harpalus will show the clypeus, the stipes and the mandibular teeth proper of the harpaloid lineage described by Brandmayr et al. (1980). Likewise, a molecular analysis will show that Baetico- harpalus will appear in a clade close to Harpalus and Pseudoophonus and separate from Ophonus and Cryptophonus.

3. Harpalus (Baeticoharpalus) lopezi Serrano & Lencina, sp. n.

Type material. Holotype male labeled with a white card: “Pico Almadén, 2,097 m. Mancha Real, Jaén. 16.4.2005. M. López leg.”, and a red Fig. 1. Habitus of Harpalus (Baetiocharpalus) lopezi label indicating HOLOTYPE. The male genitalia sp. n. The bar equals 5 mm. is mounted in a transparent plastic card on the same pin. Paratypes: two males and two females labeled very slightly arcuate. Frons with indistinct with a white card stating “Hispania. Torres (J). Si- clypeo-ocular furrows. Eyes moderately protrud- erra Mágina, Pico El Almadén. 2,032 m. 1-V- ing, neck normally strechted. Temples pubescent, 2006. Lencina, Baena, López, Bujalance & Cas- the pubescence is also present in the supraorbital tro leg.”, and a red label stating PARATYPE. The grooves, isolated setigerous punctures are also male genitalia of both males are mounted in a found in the epistome and the frons. Paraglossae transparent plastic card on their respective pin. pubescent, ligular sclerite narrow, with two long The holotype and two paratypes are deposited ventro-apical setae, first labial palpomere without in the collection of the Department of Zoology of the diagonal carina present in Ophonus.Mentum the University of Murcia. The other paratypes are without median tooth, sides not thickened, sub- deposited in the collection of José Luís Lencina. mentum with short pubescence (Fig. 2). Description.HabitusasinFig.1.Length9.0 Pronotum. Disk flattened. Anterior margin mm (holotype), 9.7–8.0 mm (paratypes). Body beaded only close to angles, posterior margin en- relatively slender, shiny black without dorsal tirely beaded. Anterior basis regularly arcuate, microsculpture. First antennomere pale reed, sec- angles moderately protruding. Posterior angles ond infuscated, 3–11 piceous but slightly paler to obtuse, well-developed. Sides moderately sinu- apex. Palpi pale reed. Legs black with tarsi pale ate before hind angles. Basal impression vaguely red (pro- and mesotarsi) or piceous (metatarsi). indicated with some coarse punctures. Sides pu- Wingless. bescent, the pubescence extended on the anterior Head. Anterior margin of labrum almost angles and on the posterior basis up to to the basal stright, not arcuate. Anterior margin of epistome impression. 196 Serrano & Lencina • ENTOMOL. FENNICA Vol. 19

Fig. 3. Male genitalia of H. (B.) lopezi sp.n.–a.Me- dian lobe of aedeagus. – b. Details of the apex in dorsal view. The bar equals 1 mm.

is well developed and is almost straight, moderately widened towards tip and without ventral serrate shaft (Fig. 3b). Endophallus with two large teeth, one medial, other apical, a basal field Fig. 2. Details of bucal parts of H. (B.) lopezi sp. n. The bar equals 1 mm. of macrotrichia and a more distal and dorsal field of microtrichia (Fig. 3a). Specific epithet. The species is named after Elytra. Relatively slender and almost parallel- Marcos López, who kindly offered us the first in- sided, slightly sinuate before apex. Humeral tooth dividual of the new subgenus and species. present. Disk flattened, intervals almost flat, Geographic distribution. The species is striae well marked but impunctate. Scutellar stria solely known from the type locality, the peak of with basal setigerous puncture. Normal seti- Almadén, located in Sierra Mágina (province of gerous puncture in the last third of the third stria Jaén, Andalusia, south Spain), coordinates or the third interval. Pubescence well developed 37.734421 (latitude), –3.523874 (longitude). in intervals 6–9, almost absent in intervals 1–5, Habitat. The five individuals of the new spe- but dense on the beginning and the apex of all in- cies were collected under stones in the meadows tervals. surrounding the Almadén Peak (Fig. 4), between Thorax (ventral sclerites) and abdomen. Pro- 1,950 and 2,023 m of altitude. The vegetation sternum pubescent, propleura sparsely punctate. corresponds to the associations that dominate the Metaepisternae longer than wider (L/W = 1.2– Supramediterranean and Oromediterranean belts 1.4) with sparse punctures and pubescence. Me- of the Betic Mountains, including the Sierra Ne- tacoxa with few isolated punctures. Last abdomi- vada (Mota et al. 2002). Common plants of the nal sternites pubescent with long hairs at each area include Erinacea anthyllis, Berberis his- side of median line. panica, Prunus ramburii, Phlomis crinita, Genis- Antennae and legs. Antennae of average size ta versicolor, Thymus serpylloides, Juniperus for Harpalus, reaching the beginning of elytra, communis ssp nana, Hormathophyla spinosum with pubescence from the 2/3 of the third an- and Sideritis glacialis. tennomere. The first antennomere has an extra The carabid assemblage of the area is proper subterminal seta before the long apical seta. Pro- of stony places located in meadows at high alti- and mesotarsi of males clearly dilated with ven- tudes of Betic sierras. Collected species were tral biseriate adhesive vestiture. All tarsomeres Leistus spinibarbis (Fabricius 1775), Notiophilus lack the dorsal pubescence existing in Ophonus. biguttatus (Fabricius 1779), Calathus baeticus Protibiae with 3–4 small spines in the outer distal mateui Nègre, 1969, Zabrus ambiguus Rambur, apex. First metatarsum shorter than second plus 1837, Carterus fulvipes Latreille, 1817, Harpalus third. honestus (Duftschmid, 1812), H. sulphuripes Male genitalia. Median lobe of aedeagus with goudotii Dejean, 1829, H. wagneri Schauberger, the typical shape of Harpalus (Fig. 3a), the apex 1926, H. serripes (Quensel 1806), Ophonus azu- ENTOMOL. FENNICA Vol. 19 • Baeticoharpalus, a new carabid subgenus 197

Fig. 4. A view of the type locality of H. (B.) lopezi sp. n., the Alma- den Peak in Sierra Mágina (province of Jaen, south Spain). reus (Fabricius, 1775), Trymosternus cordatus central Betic Mountains. This makes speciation Rambur, 1837, Cymindis etrusca baetica Ram- of cold-adapted taxa difficult during the xeric bur, 1837, C. lineola (Duftschmid, 1812), Micro- interglacial periods. It therefore seems more rea- lestes abeillei (Brisout de Barneville, 1885), sonable to hypothesise that Sierra Mágina is a rel- Philorhizus crucifer Lucas, 1846 and Brachinus ict locality of a formerly wider distribution area of pateri Puel, 1938. The new species possibly ex- H. lopezi . It is expected that a molecular analysis ists in the whole Supramediterranean belt but fur- may help in unravelling its evolutionary history. ther sampling is needed to determine its alti- tudinal range. Remarks. Adults of Harpalus (Baetico- 4. Key to Harpalus and related harpalus) lopezi are easily recognized from re- supraspecific taxa of the Iberian lated species of Harpalus, particularly from the Peninsula Iberian Peninsula, because of the unique combination of pubescence and mouth characters (see 1. Paraglossae glabrous (one or two isolated the key below). The origin of the new species is setae may be found in Cryptophonus). First uncertain. On morphological grounds there are labial palpomere with a diagonal carina. 2 no close relatives among the Iberian Harpalus. – Paraglossae pubescent, at least with 4–6 short Further, its occurrence in the central part of the lateral setae close to apex. First labial palpo- Betic Mountains does not support any clue on mere without a diagonal carina except for biogeographical grounds. These mountains (that Pseudoophonus.4 include Sierra Mágina) are poor in endemic spe- 2. Temple pubescent. Elytra uniformly punctate cies. There are three hypogean taxa (belonging to or pubescent at least in the external intervals. the genera or subgenera Typhloreicheia Hold- 3 haus, 1924, Geocharis Ehlers, 1883 and Typhlo- – Temple glabrous. Elytra glabrous except for charis Dieck, 1869) and two epigean (Tinautius the normal setigerous puncture of the third in- Mateu, 1997 and Laemostenus, Bonelli, 1810). terval; a small row of punctures may be found This fact is explained because the central Betic apicad in the seventh (and occasionally the Mountains are a transitional area between the fifth) interval, or in the eighth interval. northern carabid fauna coming from Europe and genus Cryptophonus the Sierra Nevada (Serrano et al. 2003). There are 3. Elytra with uniform dorsal pubescence. few peaks surpassing 2,000 m in altitude in the Metatarsi with dorsal pubescence (it is absent 198 Serrano & Lencina • ENTOMOL. FENNICA Vol. 19

in rare species). Metacoxae with short and middle), but easily lost in H. dispar (in that dense pubescence. genus Ophonus case the frons is pubescent). – Elytra with dorsal pubescence limited to the subgenus Artabas outer intervals, the anterior basis and the apex. Metatarsi without dorsal pubescence, only a Acknowledgements. Thanks are due to Marcos López for few, short and apical setae may be present. offering us the first individuals of the new species and to Manuel Baena, José Luís Bujalance and Alejandro Castro Metacoxae with large but sparse setae. who also collaborated in collecting more individuals of this genus Typsiharpalus species. Patricia Bermejo Gómez much helped with the 4. First labial palpomere with a diagonal carina. drawings. This work has been supported by project Metatarsi pubescent. genus Pseudoophonus 00595/PI/04, of the Fundación Seneca, Consejería de Educación de Murcia. – First labial palpomere without a diagonal carina. Metatarsi glabrous. 5 5. Prothorax with long, erect setae anteriorly. References Mentum without median tooth. genus Pangus Brandmayr, P., Ferrero, E. & Zetto-Brandmayr, T. 1980: Larval versus imaginal taxonomy and the systematic – Prothorax without long, erect setae anteriorly. status of the ground beetle taxa Harpalus and Ophonus Mentum often with a median tooth. (Coleoptera: Carabidae, Harpalini). — Entomol. Gen. 6. genus Harpalus 6 (2/4): 335–353. 6. Dorsal pubescence of elytra uniform. Habitus Brandmayr, P. & Zetto-Brandmayr, T. 1982: Identifica- zione di larve del genere Ophonus Dejean, 1821 Ophonus-like. subgenus Semiophonus (sensu novo) e note bionomiche (Coleoptera, Carabi- – Elytra glabrous or with short uniform pubes- dae). — Mem. Soc. Entomol. Ital. 60: 67–103. cence limited to outer intervals and irregular Kataev, B. M. 1987: (Ground-beetles of the genus Har- sparser punctures in inner intervals. Habitus palus Latr., group affinis). — Tr. Zool. Inst. Akad. Harpalus-like. 7 Nauk. SSSR 170: 3–41. Kataev, B. M. 1991: (A new species of the genus Harpalus 7. Temple, frons, sides of pronotum and outer of the “anxius” species group (Coleoptera, Carabidae) intervals (6–8) of elytra pubescent. Mentum from Asia Minor). — Akad. Nauk. URSS Zool. 9: without median tooth (H. dispar has frons but 147–149. not temple pubescence, sides of pronotum are Kataev, B. M. 2001: Revision of the Harpalus honestus group from the Himalaya and a review of the Harpalus not sinuate before hind angles and has species from Nepal (Coleoptera: Carabidae). — Rus- mentum tooth). subgenus Baeticoharpalus sian Entomol. J. 10 (4): 357–384. – Temple glabrous, frons also glabrous except Kataev, B. M., Wrase, D. W. & Ito, N. 2003: Subtribe for H. dispar, only sides of pronotum and Harpalina Bonelli, 1810. — In: Löbl, I. & Smetana, A. outer intervals of elytra may be pubescent. (eds.), Catalogue of Palaearctic Coleoptera. Vol. 1 Archostemata, Myxophaga, Adephaga: 367–397. Mentum with or without median tooth. 8 Apollo Books, Stenstrup. 8. Scutellar stria rudimentary and poorly delim- Lorenz, W. 2005: Systematic list of extant ground beetles ited. Elytra flattened with pubescence in all of the World. 530 pp. — Tutzing, edited by the author. intervals but sparser and irregular in inner Martínez-Navarro, E. M., Galián, J. & Serrano, J. 2005: Phylogeny and molecular evolution of the tribe ones. Mentum without median tooth. Harpalini (Coleoptera, Carabidae). inferred from mi- subgenus Licinoderus tochondrial cytochrome-oxidase I. — Mol. Phyl. Evo- – Scutellar stria well developed. Elytra vari- l. 35: 127–146. able, from flattened to convex, pubescence, Mota,J.F.,Pérez-García,F.J.,Jiménez,M.L.,Amate,J.J. when present, limited to outer intervals (ex- & Peñas, J. 2002: Phytogeographical relationships among high mountain areas in the Baetic Ranges cept in H. dispar). Mentum usually with well (South Spain). — Global Ecol. Biogeogr. 11: 497– developed median tooth. 9 504. 9. Sides of pronotum with a single large seta Noonan, G. R. 1976: Synopsis of the supra-specific taxa of close to or just in the middle. the tribe Harpalini (Coleoptera: Carabidae). — subgenus Harpalus Quaest. Entomol. 12: 3–87. Serrano, J., Lencina, J. L., & Andújar, A. 2003: Distribu- – Sides of pronotum with 6–8 long setae tion patterns of Iberian Carabidae (Insecta, Coleop- (though shorter than the normal seta in the tera). — Graellsia 59: 129–153.