Two New Species of Typhlocharis Dieck of the Gomezi Species Group from Portugal (Coleoptera: Carabidae) Author(s): Artur R. M. Serrano, Carlos A. S. Aguiar, Sónia J. R. Proença Source: The Coleopterists Bulletin, 59(2):239-249. 2005. Published By: The Coleopterists Society DOI: http://dx.doi.org/10.1649/752 URL: http://www.bioone.org/doi/full/10.1649/752
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The Coleopterists Bulletin, 59(2):239–249. 2005.
TWO NEW SPECIES OF TYPHLOCHARIS DIECK OF THE GOMEZI SPECIES GROUP FROM PORTUGAL (COLEOPTERA: CARABIDAE) ARTUR R. M. SERRANO, CARLOS A. S. AGUIAR AND SO´NIA J. R. PROENC¸A Centro de Biologia Ambiental/Departamento de Biologia Animal Faculdade de Cieˆncias da Universidade de Lisboa R. Ernesto de Vasconcelos, C2 1749-016 Lisboa, PORTUGAL
[email protected] Abstract Two endogean carabid species of Typhlocharis in the T. gomezi species group are described, Typhlocharis passosi, new species and Typhlocharis fozcoaensis, new species. This is the first record for Portugal of species belonging to the gomezi group. We provide diagnostic characters for these species, descriptions of the new species, affinities to putative relatives and a key for the identification of the gomezi species group.
Resumo Neste trabalho sa˜o descritas duas espe´cies de coleo´pteros carabı´deos endo´geos de Typhlocharis do grupo gomezi, Typhlocharis passosi, espcie nova e Typhlocharis fozcoaensis, espcie nova. Este e´ o primeiro registo de espe´cies daquele grupo para Portugal. Este estudo faculta as principais caracterı´sticas para diagnosticar as duas novas espe´cies, as suas afinidades com as formas mais pro´ximas e uma chave de identificac¸a˜o das espe´cies conhecidas do grupo gomezi.
Typhlocharis Dieck (Coleoptera: Carabidae: Trechinae: Bembidiini) is distributed throughout the Iberian Peninsula (Europe), Morocco and Tunisia (North Africa) (Jeannel 1963). This genus is considered to be very old and can be found in the lusitanian, the lionigurian, the betico-riffian and the numidian massif vestiges (Jeanne 1973). Typhlocharis belongs to the subtribe Anillina and is considered to be the richest member of the subtribe in the Iberian Peninsula with 37 known species, according to the catalogue of the ground beetles of the Iberian Peninsula (Serrano 2003). All species belonging to this genus are anophthalmous (eyes absent). They are endogean and occur in the soil, but can also be collected on the bottom surface of stones deeply buried in the soil. The majority of the species belonging to this genus have very restricted distributions, probably because of isolation by physical barriers associated with a low dispersal capacity (e.g., they cannot disperse by flight because they are wingless). During recent years the knowledge related to the systematics and distribution of Typhlocharis species present in Portugal has increased considerably (Serrano and Aguiar 2000, 2001, 2002). Nevertheless, our knowledge of this genus in this country is still incomplete. Six known species of Typhlocharis occur in Portugal, including: T. quadridentata Coiffait, T. algarvensis Coiffait, T. sarrius Serrano and Aguiar, T. singularis Serrano and Aguiar, T. elenae Serrano and Aguiar and T. gomesalvesi Serrano and Aguiar (Coiffait 1968, 1971; Serrano and Aguiar 2000, 2001, 2002). All belong to the outereloi and silvanoides species groups (sensu Zaballos and RuizTapiador 1997). Field work was conducted in several locations between Douro and Tejo rivers in Portugal that resulted in the collection of specimens belonging to two new species of the gomezi group. According to Zaballos and Ruiz-Tapiador (1997) the adults 239
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belonging to the species of the gomezi group are recognized by the following combination of characters: a) clypeus with one median tooth and b) internal apical angle of intermediate and hind tibiae dentate. The three known species of this group can be found in the Ca´ceres region in Spain (Zaballos and Farino´s 1995) (Fig. 6). This work aims to describe the two new species, provide notes about relationships with known species of the gomezi group and provide a key for the identification of the species belonging to this group. The specimens were collected using Berlese apparatus and by direct hand collecting on the bottom surface of deeply buried stones in fragments of pristine-type Mediterranean forest habitats with patches of human-induced land uses: holm-oaks and hawthorn bushes (Quercus coccifera Linnaeus and Crataegus monogyna Jacq.) (Serra d’Aire e Candeeiros) or olive trees (Olea europaea Linnaeus), rosemary bushes (Lavandula sp.) and some Cistaceae species (Vila Nova de Foz Coˆa). Typhlocharis passosi Serrano and Aguiar, new species (Figs. 1, 3a–b, 4a–b, 5a, Map Fig. 6) Diagnosis. Anophthalmous, body parallel, depressed and brownish-yellow; integument microreticulate, with scattered pubescence. Elytron with seven (4þ3) marginal umbilicate setae; apical edge with one tooth near apex of seventh stria (males and females), sutural one absent. Hind trochanters rounded in both sexes. Abdominal male sternum II with a median tubercle near the posterior margin, abdominal female sternum II with one deep posteriolateral fovea on each side. Aedeagus (Figs. 3 a–b) with median lobe sickle-shaped, basal lamina markedly arcuate; internal sac in central area with one arciform sclerite followed apically by fusiform membrane. Description. Length of holotype: 1.3 mm. Length of paratypes: 1.1–1.3 mm (males), 1.2–1.5 mm (females). Head (Figs. 1a–b) robust, more or less as long as wide [length: 0.25–0.32 mm (males) and 0.26–0.32 mm (females), width: 0.26–0.31 mm (males) and 0.25–0.32 mm (females)] with hexagonal microsculpture and slightly depressed in the middle of the frons; anterior margin of clypeus with one strong median tooth; vertex with transverse microsculpture that is arranged in parallel ridges to form a file ( pars stridens) in the area below the anterior margin of pronotum (Fig. 4a). Cephalic chaetotaxy (large setae): labrum with three pairs of setae (those on sides longer), one pair on sides of clypeus and two pairs close to frontal sulcus, a pair of supraocular setae present over each eye and two-three posterior pairs of setae between vertex and lateral carinae. Antennae moniliform and mouth-parts (Fig. 1b) with no special features; as for other members of the genus. Pronotum quadrangular (Fig. 1c), as long as wide [length: 0.30–0.37 mm (males) and 0.29–0.40 mm (females), width: 0.30–0.35 mm (males) and 0.29–0.36 mm (females)], slightly narrowed towards posterior angles which are faintly marked; disk flattened, with one central and two lateral slight sulci; anterior margin right; lateral margins with two or three minor denticles near the posterior angles; disk slightly depressed near the posterior margin, this slightly expanded near the posterior angles. Chaetotaxy: three longitudinal series of minute setae between midline and lateral margins directed anteriad and inward; one anterior seta on each side in anterior quarter, one posterior seta on hind angle; five-six pairs of setae near the anterior margin. Elytra (Fig. 1d) almost twice longer than wide [length: 0.61–0.70 mm (males) and 0.61–0.77 mm (females), width: 0.32– 0.36 mm (males) and 0.32–0.38 mm (females)], parallel and oval posteriorly, with a slightly longitudinal carinae at the beginning of seventh stria; more or less depressed on the disk; transverse scutellar organ present near the beginning of suture; scutellar region strongly punctured; humeral angles rounded, with a tooth in the beginning of carinae; lateral margins serrate, teeth decreasing in size posteriorly; apical margin (Fig. 1e) with one tooth coinciding with end of seventh stria, without tooth at sutural angles. Chaetotaxy: part of the pubescence of the disk arranged in four lines, these short setae are erect and slightly directed anteriad; umbilicate series with four setae in the front group and three in the hind group (4 þ 3) (Fig. 1e). Legs with robust femora, tibiae dilated distally in both sexes; median and hind tibiae of males with two or three teeth at internal apical angle, much more developed in the latter (Fig. 4b); trochanters of third pair round, similar in both sexes (Figs. 1g–h), protarsus with segments not dilated. Abdominal sternum
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Fig. 1. Typhlocharis passosi. a) Head (dorsal view); b) head and anterior part of pronotum, (ventral view); c) pronotum (dorsal view); d) elytra (dorsal view), e) right elytron (latero-dorsal view); f) elytra (apical view); g) thorax (part) and abdomen (male, ventral view); and h) thorax (part) and abdomen (female, ventral view). II of males with one median tubercle near the posterior margin (Fig. 1g), females with one deep posteriolateral fovea on each side (Fig. 1h). Male genitalia (Figs. 3a–b) in lateral aspect (Fig. 3a) with median lobe sickle-shaped, basal lamina markedly arcuate; median lobe in dorsal aspect (Fig. 3b) with apex more or less thin, bent to the left; internal sac in central area with one arciform
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Fig. 2. Typhlocharis fozcoaensis. a) Head (dorsal view); b) head (ventral view); c) pronotum (dorsal view); d) elytra (dorsal view); e) left elytron (latero-dorsal view); f) elytra (apical view); g) thorax (part) and abdomen (male, ventral view); and h) thorax (part) and abdomen (female, ventral view). sclerite followed apically by fusiform membrane (Fig. 3a); parameres bisetulose apically, lateral aspect of left paramere as in Fig. 3a. Female genitalia (Fig. 5a) with ovipositor gonocoxites weakly sclerotized, each one in ventral aspect with one apical seta and one medium seta; internal genital tract with spermathecal duct short and spermatheca spheroid.
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Fig. 3. Aedeagus, Typhlocharis passosi. a) Median lobe and left paramere (lateral view); b) median lobe (dorsal view); Typhlocharis fozcoaensis. c) median lobe and left paramere (lateral view); d) median lobe (dorsal view).
Type Series. Holotype #: Portugal, Alvados (Serra d’Aire e Candeeiros) (UTM: 29SND1976), 30.I.2002. Paratypes: 22 ## and 38 $$ (2 ## and 2 $$ gold coated), same locality and date; Carvalhal (UTM: 29SND0677), 2 ## and 5 $$ (1 # and 2 $$ gold coated), 6.V.2003; Souro˜es (Serra d’Aire e Candeeiros) (UTM: 29SND1065), 5 ## and 6 $$; Teira (Serra d’Aire e Candeeiros) (UTM: 29SND0661), 6 ## and 3 $$. Holotype and paratypes deposited in the collection of the senior author, Department of Animal Biology (Lisbon).
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Fig. 4. Typhlocharis passosi. a) Stridulatory organ (pars stridens) (dorsal view); b) hind tibia of male (ventral view) with latero-apical tooth (arrow). Typhlocharis fozcoaensis. c) stridulatory organ (pars stridens) (dorsal view); d) hind tibia of male (ventral view) with latero-apical tooth (arrow).
Etymology. This new species is dedicated in modest homage to the memory of the late Eng8 Passos de Carvalho, an eminent Portuguese applied entomologist (agriculture) from Estac¸a˜o Agrono´mica Nacional de Oeiras, who was a great friend and a reference for the senior author. Affinities. The new species belongs to the gomezi group based on the presence of apical teeth of the second and third tibiae (males) and a median tooth of the clypeus. However, it is differentiated within the gomezi group by the presence of a median tubercle in sternum II (males) near the posterior margin (Fig. 1g). Some of the other species belonging to Typhlocharis also present this interesting type of sexual dimorphism (e.g., T. monasticus Zaballos and Wrase, T. peregrinus Zaballos and Wrase, and T. navaricus Zaballos and Wrase) (Zaballos and Wrase 1998). However, the phylogenetic importance of this structure is uncertain because it also occurs in species of the monasticus and outereloi groups (sensu Zaballos and Ruiz-Tapiador 1996; Zaballos and Wrase 1998). Within the gomezi species group, only T. wrasei Zaballos and Farino´s and the new species present a similar form of the median lobe of the aedeagus in lateral view. Differences in the conformation of the apex of the median lobe, left paramere and internal sac separate T. passosi from all of the other species belonging to this group. The pattern of the umbilicate series of the elytra (4 þ 3) found in the new species is only similar to the one found in T. gomezi Zaballos. All the other known species of the gomezi group present a pattern of 4 þ 2, including T. fozcoaensis n. sp. (see next description). This fact suggests that small species of Typhlocharis, such as those belonging to this group, manifest a tendency to diminish the number of setae within the posterior group of the umbilicate series. The rounded-shape of the hind trochanters of the new species and the number of foveae (one) present in abdominal sternum II are also found in T. gomezi. The spermatheca of Typhlocharis passosi has
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Fig. 5. Female genitalia. a) Typhlocharis passosi; and b) Typhlocharis fozcoaensis.
a similar form (spheroid) to that found on the other members of the gomezi group, but the number of apical gonocoxite setae is different (one vs. two). With the data available at the moment it is very difficult to determine the real affinities of T. passosi. Taking into account the analyzed characters, the new species seems to have a close relationship with T. wrasei and T. gomezi.
Typhlocharis fozcoaensis Serrano and Aguiar, new species (Figs. 2, 3c–d, 4c–d, 5b, Map Fig. 6) Diagnosis. Anophthalmous, body parallel, depressed and brown; integument microreticulate, with scattered pubescence. Elytron with six (4 þ 2) marginal umbilicate setae; apical edge with two pairs of teeth, one sutural and other at the apex of the seventh stria (males and females). Hind trochanters without sexual dimorphism. Abdominal male sternum II without any special structure, abdominal female sterna II and III with one posteriolateral fovea and one anterolateral fovea in each side, respectively. Aedeagus
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Fig. 6. Geographic distribution of the genus Typhlocharis (ellipses) and of the five species of the gomezi group. T. passosi (P); T. fozcoaensis (F); T. gomezi (G); T. hiekei (H); T. wrasei (W).
(Figs. 3c–d) with median lobe sickle-shaped, but the apex erect, basal lamina markedly arcuate; internal sac in central area with one circled sclerite. Description. Length of holotype: 1.4 mm. Length of paratypes: 1.3–1.4 mm (males and females). Head (Fig. 2a–b) as long as wide [length: 0.27–0.30 mm (males) and 0.26–0.29 (females), width: 0.29–0.30 mm (males) and 0.29–0.30 mm (females)] with hexagonal microsculpture and faintly depressed in the middle of the frons; clypeus with one strong median tooth in anterior margin; vertex with transversal microsculpture, forming a file of parallel ridges ( pars stridens) in the area bellow the anterior margin of pronotum (Fig. 4c). Cephalic chaetotaxy (large setae): labrum with three pairs of setae (those on sides longer), one pair on sides of clypeus and two pairs close to frontal sulcus, two pairs of supraocular setae (anterior and posterior) and two-three pairs of setae on the posterior region between vertex and lateral carinae. Antennae
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moniliform and mouth-parts (Fig. 2b) with no special features, as for other members of the genus. Pronotum quadrangular (Fig. 2c), slightly longer (1.1–1.2 times) than wide [length: 0.37–0.39 mm (males) and 0.37–0.38 (females), width: 0.31–0.34 mm (males) and 0.32–0.34 mm (females)], narrowed towards posterior angles which are slightly marked; disk depressed, with longitudinal sulci, one central and two lateral; anterior and posterior margins straights; lateral margins subparallel with two or three minor denticles near the posterior angles; disk depressed near the posterior margin. Chaetotaxy: three longitudinal series of minute setae between midline and lateral margins directed anteriad; one anterior seta on each side in anterior quarter, one posterior seta on hind angle; one seta near the anterior angles; four pairs of setae near the anterior margin. Elytra (Fig. 2d) twice or even more longer than wide [length: 0.64–0.68 mm (males) and 0.66–0.70 mm (females), width: 0.30–0.34 mm (males) and 0.34–0.35 mm (females)], parallel and oval posteriorly, with a subtle longitudinal carinae at the beginning of seventh stria; flattened on the disk; transverse scutellar organ present near the base of suture; scutellar region strongly punctured; humeral angles well marked and rounded, with a minor tooth in the base of carinae; lateral margins serrate, teeth decreasing in size posteriorly; apical margin (Fig. 2f) dentate in the sutural angles and with one tooth coinciding with end of seventh stria (males and females). Chaetotaxy: part of the pubescence of the disk is arranged in three lines, these setae are erect and slightly directed anteriad; umbilicate series with four setae in the front group and two setae in the hind group (4 þ 2) (Fig. 2e). Legs with robust femora, tibiae dilated distally in both sexes; median and hind tibiae of males with one or two teeth at internal apical angle, much more developed in the latter (Fig. 4d); trochanters of third pair round-shaped, similar in both sexes (Figs. 2g–h), protarsus with segments not dilated. Abdominal sternum II of males without a median tubercle near the posterior margin (Fig. 2g); females with deep foveae, one posteriolateral pair in the abdominal sternum II and one anterolateral pair in the sternum III, respectively (Fig. 2h). Male genitalia (Figs. 3c–d) in lateral aspect (Fig. 3c) with median lobe sickle-shaped, but with apex erect, basal lamina markedly arcuate; median lobe in dorsal aspect (Fig. 3d) with apex largely rounded, bent to the left; internal sac in central area with one circular sclerite (Fig. 3a); parameres bisetulose apically, lateral aspect of left paramere as in Fig. 3c. Female genitalia (Fig. 5b) with ovipositor gonocoxites weakly sclerotized, each one in ventral aspect with one apical seta and one medium seta; internal genital tract with spermathecal duct short and spermatheca spheroid, spermathecal gland long, with proximal region membranous and apical region more or less sclerotized.
Type Series. Holotype #: Portugal, Vila Nova de Foz Coˆa (UTM: 29TPF5343), 5.III.2003. Paratypes: 2 ## and 3 $$ (1 # and 2 $$ gold coated), same locality and date. Holotype and paratypes deposited in the collection of the senior author, Department of Animal Biology (Lisbon). Etymology. The name of the new species is the latinized adjectival from the composition of the last two names of the village (Vila Nova de Foz Coˆa) where the specimens have been found. Affinities. This new species also belongs to the gomezi group. It is well differentiated from the other species of this group by the presence of one anterolateral foveae on each side of the abdominal female sternum III (Fig. 2h). Like the other species belonging to the gomezi group with the exception of T. passosi, the abdominal male sternum II of this species does not present a median tubercle near the posterior margin. Within this group, only T. gomezi Zaballos presents an aedeagus with a similar form of the internal sac of the median lobe in lateral view. Particular features related with the forms of the median lobe, the left paramere and the internal sac easily separate the new species from all the others of the same group. The pattern of umbilicate series of elytra (4þ2) is similar to the one found on T. wrasei Zaballos and Farino´s and T. hiekei Zaballos and Farino´s. The rounded-shape of the hind trochanters of the new species cluster the species T. fozcoaensis, T. gomezi and T. passosi together. The new species, as well as T. gomezi and T. hiekei, presents an identical number of foveae in the abdominal sternum II of the females (one pair). Typhlocharis fozcoaensis presents a spermatheca with a similar form (spheroid), to the ones that can be found in the other members of the gomezi group, but the number of apical gonocoxite setae is different
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(one vs. two). In this aspect both new species are identical. Taking into account the comparison of all analysed characters, the new species seems to have a closer relationship with T. gomezi. As a final remark, we point out that all species of the gomezi group found until now were collected in the lusitanian massif of the Iberian Peninsula (three species in Spain and two in Portugal) (Zaballos 1991; Zaballos and Farino´s 1995; Serrano and Aguiar, this work) (Fig. 6). However, while the Spanish assemblage is within the same region (Ca´ceres), the greatest distance found among species being 55 km (between T. gomezi and T. hiekei) (Zaballos and Farino´s 1995), the two Portuguese species are separeted by a greater distance (around 200 km from each other, an equal distance existing between each new species and the Spanish assemblage). Key to Species of the Typhlocharis gomezi Species Group The gomezi species group includes the following five species: T. gomezi, T. wrasei, T. hiekei, T. passosi and T. fozcoaensis. All the species are distinguished from one another by character states in the following key: 1. Abdominal sternum II of male with a median tubercle near the posterior margin, female with one deep pair of posteriolateral foveae. Umbilicate series 4 þ 3. Middle lobe of aedeagus as in Figures 3a–b - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - T. passosi 19. Abdominal sternum II of male without a median tubercle near the posterior margin, female with one or two pairs of lateral foveae - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 2 2. Trochanter and internal edge of femur of third pair of legs dentate in male. Umbilicate series 4 þ 2 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 3 29. Trochanter and internal edge of femur of third pair of legs not dentate in male. Umbilicate series 4 þ 2 or 4 þ 3 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 4 3. Trochanter of second pair of legs dentate in male, rounded in female. Internal edge of second pair of femora rounded in both sexes. Abdominal sternum II of female with one pair of shallow foveae at hind angles. Middle lobe of aedeagus as in Figure 3 of Zaballos and Farino´s (1995) - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - T. hiekei 39. Trochanter and internal edge of femur of second pair of legs dentate in male, rounded or angular in female. Abdominal sternum II of female with two pairs of lateral foveae (one front pair and one hind pair). Middle lobe of aedeagus as in Figure 2 of Zaballos and Farino´s (1995) - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - T. wrasei 4. Umbilicate series 4 þ 2. Abdominal sternum III of females with one pair of anterolateral foveae. Middle lobe of aedeagus as in Figures. 3c–d - - - - - - - - - T. fozcoaensis 49. Umbilicate series 4 þ 3. Abdominal sternum III of females without any lateral foveae. Middle lobe of aedeagus as in Figure 1 of Zaballos and Farino´s (1995) - - - T. gomezi
Acknowledgments We are grateful to Luisa Ganc¸o and Telmo Antunes for photographic assistance. We are indebted to two anonymous reviewers for their helpful comments and suggestions that improved the manuscript. Carla Rego and Maria Jose´ Boavida helped us to improve the English version. This work was partially financed by Centro de Biologia Ambiental (CBA). Literature Cited Coiffait, H. 1968. Nouveaux Anillini du Maroc et du Sud de la Peninsule Iberique. Bulletin de la Socie´te´ des Sciences Naturelles et Physiques du Maroc 48(3–4):55–66.
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Coiffait, H. 1971. Contribution a la connaissance du genre Typhlocharis (Col. Carabidae). Description d’une espe`ce nouvelle du Portugal. Annales de Spe´le´ologie 26(2):463–467. Jeanne, C. 1973. Sur la Classification des Bembidiides endoge´s de la Re´gion EuroMe´diterrane´enne (Col. Carabidae, Bembidiinae, Anillini). Nouvelle Revue d’Entomologie 3(2):83–102. Jeannel, R. 1963. Monographie des «Anillini», Bembidiides endoge´s (Coleoptera Trechidae). Me´moires du Muse´um National D’Histoire Naturelle (N.S.), Se´rie A, Zoologie 28(2): 33–204. Serrano, A. R. M., and C. A. S. Aguiar. 2000. Description of two new endogean beetle species (Coleoptera, Carabidae) from Portugal. Boletim da Sociedade Portuguesa de Entomologia 7(13):149–158. Serrano, A. R. M., and C. A. S. Aguiar. 2001. Three new endogean beetles (Coleoptera, Carabidae) from Portugal. The Coleopterists Bulletin 55:172–180. Serrano, A. R. M., and C. A. S. Aguiar. 2002. The genus Typhlocharis Dieck, 1869 (Coleoptera: Carabidae) in Portugal: description of two new species and faunistic notes. Boletim da Sociedade Portuguesa de Entomologia 7(16):181–197. Serrano, J. 2003. Cata´logo de los Carabidae (Coleoptera) de la Penı´nsula Ibe´rica. Monografias S.E.A 9:1–130. Zaballos, J. P. 1991. Dos nuevos Typhlocharis de Extremadura (Espan˜a) (Coleoptera, Trechidae). Nouvelle Revue d’Entomologie (N.S.) 8:331–336. Zaballos, J. P., and G. P. Farins. 1995. Systematics of the genus Typhlocharis Dieck: the T. gomezi species Group (Coleoptera: Caraboidea: Trechidae). The Coleopterists Bulletin 49:89–95. Zaballos, J. P., and I. Ruiz-Tapiador. 1997. Nuevos Typhlocharis Dieck (Coleoptera, Caraboidea, Trechidae) de Espan˜a. Graellsia 52:95–106. Zaballos, J. P., and D. W. Wrase. 1998. Tres nuevos Typhlocharis Dieck, 1869 (Coleoptera, Caraboidea, Trechidae) de Navarra (Espan˜a). Graellsia 54:43–52. (Received 12 May 2004; accepted 19 February 2005. Publication date 1 August 2005.)
