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THE SPECIES OF Notiobia PERTY (COLEOPTERA: CARABIDAE: HARPALINI) FROM BRAZIL

— The new species Notiobia glabrata, N. maxima and N. pseudolimbipennis are described. A key to the 11 Notiobia (s.str.) species known from Brazil, data about the synonym of N. nebrioides Perty, 1830, and Notiobia Bates, different species groups, each distributed from Brazil over the North-Western part of South Central America Mexico.


INTRODUCTION
The genus Notiobia Petty is part of the Anisodactylina, a subtribe of the large carabid tribe Harpalini. Noonan (1973) placed in Notiobia four groups previously regarded as separate genera: Diatypus Murray from the tropical region of Africa; Diaphoromerus Chaudoir from Australia, New Zealand, New Guinea, New Caledonia, the Moluccas, Timor, and Hawaii; Anisotarsus Chaudoir from North, Central, and temperate areas of South America; and Notiobia (s.str.) from Central America and tropical areas of South America.
The Neotropical species of the subgenus Anisotarsus were revised by Noonan (1981). Only two species are reported to occur in the dry areas of Brazil. The Mexican species of Notiobia (s.str.) were already revised in Noonans (1973) classification of Anisodactylina. Much less is known about the taxonomy and distribution of Notiobia (s.str.) in areas south of Mexico.
The idea to describe and key out the Brazilian species of Notiobia (s.str.) was born during an ecological study. Contrary to the rather strong taxonomicai confusion of species in the mountain areas of Panama, Colombia, and Ecuador, the taxonomy of Brazilian species is quite clear. The present paper provides accurate taxonomic names for current ecological studies and provides information useful for future revision of all Notiobia species.
As far as known, Notiobia species live in forests. Adults of all species can fly. They are spermatophagous and their larvae seem to develop only on specific fruits. Therefore, fruit fall events in the forests which are restricted in time and space cause complicated ecological adaptations of Notiobia species (Paarmann, pers. comm.). The results on the phylogenetic and zoogeographic relations of these species will help to understand the process of the ecological adaptations.

MATERIAL
Beside the vast material collected during an ecological research project by W. Paarmann, J. Adis and co-workers, and the author between 1992 and 1996, specimens of following institute collections were considered: CAS -Dept. of Entomology, California Academy of Sciences (D. Kavanaugh The terminology of the descriptions and key follows Noonan (1981Noonan ( ,1991.
Head: Labrum straight to emarginate anteriorly; clypeus broadly emarginate; mentum with prominent median tooth, mentum and submentum completely separated by a transverse suture; paraglossa slightly longer than ligula. Frons usually with moderate to prominent fovea bearing a clypeo-ocular prolongation. Eyes mostly large and protruding; width of narrowest part of gena less than maximum width of antennomere I; supraantennal ridges divergent anteriorly.
Thorax: Pronotum arcuate anteriorly, convergent posteriorly; posterior angles rounded or distinct, base lobed in most species; basal fovea more or less distinct; lateral bead complete, basal and anterior bead at least laterally distinct.
Legs: Anterior tibia with apical spur lanceolate; posterior femur usually with 2 long setae on posterior margin; posterior tarsus with segment I shorter than II+III; anterior and middle tarsi of males moderately to very strong enlarged.
Elytra: Interval III with a setigerous puncture in apical third; interval VII with a small setigerous puncture near apex and a slightly more proximal ocellate puncture. Hind wings fully developed in most specimens.
Abdomen : Sternum VI of males with a pair of ambulatory setae. Median lobe of aedeagus symmetrical, without apical disc. Valvifer flat to slightly convex, lateral margin semi-membranous and without distinct boundary. Body length: 11-13 mm. Color: Dorsum with labrum rufopiceous and clypeus yellow-piceous; remaining parts of head, pronotum, and elytra greenish, bluegreen or cupreous with metallic lustre; ventral part of body generally dark piceous; legs rufopiceous to dark rufopiceous, palpi and antenna testaceous.
Thorax: Pronotum with sides arcuate anteriorly, convergent and rectilinear posteriorly; posterior angles arcuate to slightly obtuse, base lobed; lateral depression complete, widened posteriorly and ending in the wide, shallow basal fovea; lateral bead complete, basal and anterior beads only distinct at the sides, absent in the middle. Microsculpture of very fine transverse meshes, micropunctures present.
Legs: Dorsa of all tarsi glabrous except the anterior tarsi I-IV of males which bear single small hairs. Anterior and median tarsi of males strongly expanded laterally.
Abdomen: Sternum VI of females evenly rounded apically (Fig. 4). Aedeagus with median lobe slender with pointed apex (Figs. 5,6). Everted internal sac (Fig. 20) with a large field of macrotrichia on right side. Distribution: The species is known from the type locality and from Panama, Cerro Campana.
Derivation of species name: The name expresses the similarity of the new species to N. viridula (=N. limbipennis).
Deposition of type material: The holotype and part of the paratypes are deposited at the Systematic Entomology collection of the EMPA (Manaus, Brazil). The rest of the paratypes are at the Zoological Museum of the University São Paulo, in the collection W. Paarmann and the collection of the author.
Discussion: N. pseudolimbipennis is closely related to N. viridula, a widespread species in Central and South America. However, N. pseudolimbipennis is clearly distinguished from N. viridula by the lack of the apical spine of sternum VI in females. Furthermore, the granulate lateral area of the elytra is green or green-bronze with a metallic lustre like the rest of the elytral intervals in the Brazilian specimens of N. pseudolimbipennis. It is distinctly lighter, yellow-testaceous in N. viridula. N. viridula is not known from the Amazonian lowlands. Probably N. pseudolimbipennis replaces N. viridula in this region.
Specimens without an apical spine on sternum VI of females but with bicolored elytra were found in Panama (Cerro Campana, coll. NMNH). These specimens have a green-bronze dorsal surface, but the granulate area is not extended like in N. viridula. Therefore these Panamanian specimens belong to N. pseudolimbipennis but represent a transitional form which is similar to N. viridula. Color: Dorsum with labrum and clypeus piceous; remaining parts of head, pronotum, and elytra green or cupreous with metallic lustre; ventral part of body generally dark piceous; legs piceous, palpi and antenna yellow-piceous.
Thorax: Pronotum ( Fig. 23) with sides arcuate anteriorly, convergent and rectilinear posteriorly; posterior angles distinct, not rounded, slightly obtuse, base lobed; lateral depression complete; basal fovea shallow; basal and lateral beads complete; anterior bead distinct only at the sides, obsolete in the middle. Microsculpture not distinct, surface shining; micropunctures present.
Legs: Dorsa of anterior tarsi I-IV of males pubescent, rest of tarsi of males and tarsi of females glabrous dorsally; anterior and median tarsi of males comparably slightly extended.
Distribution: The species is known from the type locality and from Peru, prov. Loreto, Rio Napo, Rio Amazonas, prov. Madre de Dios, Pakitza, Guyana, and Bolivia, prov. La Paz (one specimen each in NMNH and MPM).
Derivation of species name: The name is related to the dorsal surface of the body which lacks a microsculpture.
Deposition of type material: The holotype and part of the paratypes are deposited at the Systematic Entomology collection of the INPA (Manaus, Brazil). The rest of the paratypes are at the Zoological Museum of the University São Paulo, in the collection W. Paarmann and the collection of the author.
Discussion: N. glabrata is morphologically a transitional form between N. leiroides and N. maxima. N. glabrata differs from N. maxima by the structure of the aedeagus and the generally smaller size. N. glabrata is distinguished from N. leiroides by the distinctly slender anterior tarsi of males and the absence of microsculpture. Moreover, the typical N. leiroides lacks a prolongation of the frontal foveae to the eyes. However there are known specimens of the leiroides-growp from Peru with a clypeo-ocular prolongation of the frontal fovea and with flattened microsculpture, which are close to N. glabrata. Further studies are needed to decide if these specimens fall within the variation of N. leiroides or represent a separate species. A rather strong radiation of this groups occurs also in Panama. A revision of the Panamanian forms is needed. Head: Labrum slightly emarginate anteriorly, clypeus broadly emarginate anteriorly; frons with foveae punctiform, bearing a distinct clypeoocular prolongation to eyes; eyes large and protruding. Microsculpture not distinct, micropunctures present.
Thorax: Pronotum ( Fig. 23) with sides arcuate anteriorly, convergent and rectilinear posteriorly; posterior angles arcuate to slightly obtuse, base lobed; lateral depression complete, ending in shallow basal fovea; basal and lateral beads complete; anterior bead distinct only at sides, obsolete in middle. Microsculpture not distinct, surface shining; micropunctures present.
Legs: Dorsa of anterior tarsi I-IV in males with single fine hairs, rest of tarsi in males and tarsi in females gla-brous dorsally; anterior and median tarsi of males usually expanded laterally.
Distribution: The species is only known from the type locality.
Derivation of species name: The name refers to the body size, it is the largest kncwn species of Notiobia (s.str.).
Deposition of type material: The holotype and part of the paratypes are deposited at the Systematic Entomology collection of the INPA (Manaus, Brazil). The rest of the paratypes are at the Zoological Museum of the University São Paulo, in the collection W. Paarmann and the collection of the author. viridella Csiki, 1929: 1047 (synonym of viridula (Dejean).

DISCUSSION
The 11 species known from Brazil can be tentatively placed in 3 species groups.
The first group is characterized by granulate lateral areas of the elytra, which are in part light yellow-testaceous. This character is absent in Anisotarsus and related genera and can be considered as apomorphic. The group includes the species Notiobia disparilis, N. nebrioides, N pseudolimbipennis, N viridula, and probably N flavicinctus. All these species occur on fallen fruits of Ficus. The larval development of N. pseudolimbipennis and N. flavicinctus was also observed on fallen fruits of Ficus. However, the N. nebrioides population of Manaus (Brazil) was found to develop on fallen fruits of Vismia and Coussapoa {Paarmann, pers. comm.), and one of the populations in Loreto Boca (Peru) was collected on fallen fruits of Cecropia (specimens in NMNH). TV. nebrioides is the species with the weakest development of the granulate microsculpture.
The second group has flat elytral intervals, the microsculpture is uniform or absent. These characters are probably plesiomorphic (see also Noonan 1973). The phylogenetic relationship of the species is therefore not evident. The group includes N. áulica, N. glabrata, N. jucunda, N maxima, and N wilkensi. The larval development of these species was observed on fallen fruits of Melastomataceae or (in Peru) Cecropia. The adults also appear on fallen fruits of Ficus, where they only eat but do not reproduce (Paarmann, pers. comm.).
The third group comprises only ./V. umbrifera, which cannot be placed in one of the previous groups. This species has a spatula-shaped prolongation of the median lobe of the aedeagus, which is doubtless an apomorphic character state. Adults of N. umbrifera were frequently found on Ficus, Cecropia and on Miconia fruit fall. The larval development was only observed on Miconia fruit fall (Paarmann, pers. comm.).
All three species groups are widespread and occur in the whole area of Notiobia (s.str.) from Brazil to Mexico.

KEYS TO THE SUBGENERA AND SPECIES
Key to the subgenera Notiobia (s.str.) Perty and Anisotarsus

Key to the Notiobia (s.str.) species from Brazil
1 Postero-lateral intervals of elytra with a region of distinct granulate microsculpture, extended in some females to widest part of the elytra and/or light yellowish; subapical sinuation of elytra prominent ( Fig. 1) or not (Fig. 2) 2 -Lateral intervals of elytra without granulate microsculpture; microsculpture of all intervals uniform or lacking; lateral parts of elytra not yellowish and not lighter than the green colored inner intervals; subapical sinuation of elytra not prominent (Fig-2) 7 2(1) Subapical sinuation of elytra prominent (Fig. 1) 3 -Subapical sinuation of elytra not prominent (Fig. 2) 5 3(2) Intervals of elytra the same in males and females, not convex; median lobe of aedeagus various; dorsum green, bronze or blue-green, lateral intervals in part pale yellow-testaceous 4 -Males with intervals of elytra convex, dorsum of males bicolored in most specimens, head and pronotum golden green-cupreous, elytra black with purple tinge; median lobe of aedeagus with short but wide apex (Fig. 7). Females with convex intervals 2,4 and 6, but flat intervals 1,3,5, and 7; dorsum of females uniformly dark colored N. disparilis Bates 4(3) Sternum VI of females produced into a ventrally projected spine (Fig.  3); median lobe of male aedeagus with short but wide apex (Fig. 12); elytra distinctly bicolored, with lateral intervals granulate, pale yellow-testaceous and inner intervals green-cupreous; in females granulated area in the anterior part enlarged, covering the full anterior part of elytra N. viridula (Dejean) -Sternum VI of females rounded apically (Fig. 4); median lobe of male aedeagus with longer and more narrow apex (Figs. 5, 6); elytra not distinctly bicolored in most specimens; granulated area of elytra anteriorly not expanded to the middle part, dorsal surface more shining, green, cupreous or black with greenish, bluish or purple lustre N. pseudolimbipennis sp. n. 5(2) Body length 11-13 mm; lateral intervals of elytra of the same color as inner intervals, not pale yellow-testaceous; median lobe of aedeagus pointed (Fig. 8)..
N. nebrioides Perty -Body length 7-10 mm; lateral intervals, in some females whole elytra except a preapical macula, granulated and pale yellow-testaceous, inner intervals green or cupreous; aedeagus not as in Fig. 8 6 6 (5) Pale region of lateral intervals in both sexes narrow, sometimes indistinct; body length usually 7-8 mm; apex of median lobe of aedeagus elongated, spatula-shaped (Fig. 9,10), anterior tarsi of males much narrower than in the following species N. umbrifera Bates -Pale region of lateral intervals strongly enlarged in females, in specimens from the Andes and Central America covering the whole elytra except a subapical region, in Amazonian specimens extended to the lateral 4-5 intervals in the posterior part of elytra; median lobe of aedeagus very slender, with nearly rounded apex (Fig. 11); anterior tarsi of males very wide.