Review of the species of Paratenetus Spinola inhabiting America, north of Mexico (Coleoptera, Tenebrionidae)

Abstract The North American (north of Mexico) species of the tenebrionid genus Paratenetus Spinola are reviewed and a key is presented for their identification. Five species are recognized, P. gibbipennis Motschulsky, P. fuscus LeConte, P. punctatus Spinola and two sp. n., P. exutus [type locality: Tabusintac, Nova Scotia] and P. texanus [type locality: Port Isabel, Cameron County, Texas]. Two syn. n. are proposed: P. cribratus Motschulsky, 1868 with P. gibbipennis Motschulsky, 1868 and P. crinitus Fall, 1907 with P. fuscus LeConte, 1850. A lectotype is selected for Paratenetus punctatus Spinola. A type species is designated for Storthephora Mäklin, 1875 (Storthephora denticollis Mäklin, 1875).


Introduction
The genus Paratenetus was proposed by Spinola in 1844 for two species, P. lebasi from Colombia and P. punctatus from the United States of America. The last mentioned species was represented by three specimens originating from the collection of Baron Dejean who received them from John Eatton LeConte. Subsequently, the genus received very little attention. In North America, John Lawrence LeConte described a new species in 1850 which he obtained during his trips to Lake Superior. In 1853-54, Victor de Motschulsky, a Russian Imperial Army Colonel, made a 10-month trip to the United States and Panama and collected at several locations including New York, Niagara Falls, Cleveland, Cincinnati, Lexington, Louisville, New Orleans, Mobile, Atlanta, Washington D.C., and Philadelphia. He described two species of Paratenetus in 1868 from the material he collected in Georgia. In 1907, Fall described a new species from New Mexico. Subsequently, the genus and some of its species were briefly cited in monographic works such as Blatchley (1910), Downie and Arnett (1996) and Aalbu et al. (2002a, b).
The purpose of this paper is to review the American species occurring north of Mexico and provide a key for their identification.

Material
The study is based on the examination of about 3110 specimens borrowed from the following collections: placed the genera of Heterotarsini (except Heterotarsus) in the lagriine subtribe Lupropina of the tribe Adeliini. Parsons (1976: 211) listed Paratenetus in the lagriid subfamily Lupropinae. Doyen and Tschinkel (1982: 183) indicated that the genus may belong to the belopines, currently a valid lagriine tribe. Campbell (1991: 261) listed Paratenetus in the lagriid subfamily Goniaderinae and Aalbu et al. (2002a: 509;2002b: 484) retained also the genus in the lagriine tribe Goniaderini. On the other hand, Ferrer and Ødegaard (2005: 648) included it in the lagriine tribe Lupropini following Ardoin (1961) and Parsons (1976).
We did not investigate the taxonomic position of the genus Paratenetus but we accept, following Aalbu et al. (2002a;2002b), its placement in the tribe Goniaderini of the subfamily Lagriinae within the Tenebrionidae.
Biology. The biology of members of Paratenetus is poorly known. Many of the specimens seen in this study were collected in leaf litter in forested areas or in nests of the tent caterpillar genus Malacosoma (Lepidoptera: Lasiocampidae). All three winged species have been collected at black light. Steiner (1995: 508) commented that Paratenetus species pupate on the inner surfaces of rolled dead leaves (in which the larvae live) either hanging on fallen tree branches or on the ground.
Notes. There are two types of setae on the elytra of Paratenetus: erect and slanting. The slanting setae are characterized as subdepressed when the angle between the base of the seta and the elytra is between 10 and 40°, semierect when the angle is between 40 and 60°, and suberect when the angle is between 60 and 80°. The specimen is intact although many of the setae on the pronotum and elytra are gone. The provenance of the specimen is doubtful. In the key to the Paratenetus in his collection, Motschulsky (1868: 193) mentioned that the species was collected in Atlanta but Mobile is listed on one of the type labels. Motschulsky collected in both localities during his 10-month trip to America in 1853-54. Motschulsky's collection contains a single specimen, a male, under the name P. cribratus. It bears the following labels: "[green round disc] / Atlanta [handwritten] / type [handwritten] / Paratenetus cribratus Motch Am. bor. Atlanta [handwritten on a rectangular green label]." The specimen is missing the left antennomeres 3-11 and the posterior legs.

Key to North American (north of Mexico) species of Paratenetus
Note about synonymy. Motschulsky (1868) separated P. gibbipennis and P. cribratus on the account that the first species has the lateral denticles of the pronotum very short while the second species has strong denticles. From an examination of the types, we cannot sustain Motschulsky's affirmation; the denticles are basically of the same size on both specimens.
Diagnosis. This species and P. fuscus differ from the other three species treated by having the metaventrite very short. Paratenetus gibbipennis differs from P. fuscus by having few short erect setae on the elytra.
Description. Body dorsally reddish yellow to dark reddish brown, legs paler, yellow to reddish yellow; antennal club not darkened in most specimens; metaventrite not darker than first two abdominal ventrites. Antennomere 8 subquadrate or very slight-ly transverse. Pronotum with maximum width near midlength or slightly anterior to midlength; punctures moderately dense, not subcontiguous even over lateral half. Elytra very convex; slanting setae subdepressed, erect setae very few, short. Metaventrite short, length along midline clearly shorter than length of abdominal ventrite 2 along midline. Male protibia with calcar near middle along ventral surface; male mesotibia with short, in some specimens very short, more or less perpendicular preapical protuberance. Parameres with sides more or less parallel towards apex, apex not particularly acute (Fig. 5).
Length: 2.5-3.2 mm. Distribution. This species ranges from southern Maine to southwestern Manitoba, south to central Texas, southwestern Alabama, and central South Carolina (Fig. 10).
Labels on specimens read "in leaf litter" (6 specimens); "in leaf litter of black birch and shrubs around and on areas of exposed rock" (71); "forest litter sifting" (2); "forest litter" (3); "moist forest berlese" (1). Note about synonymy. Fall (1907: 253) described his P. crinitus and mentioned that "in crinitus the metasternum is almost as short as in fuscus, which species is, however, very distinct by its subinflated elytra, more rounded sides of the prothorax and absence of erect hairs on the upper surface." Obviously Fall did not study the syntype in LeConte's collection since the specimen bears many erect hairs. LeConte never mentioned that character in his description and obviously Fall misidentified LeConte's species. We have studied the type specimens of both species and find no structural differences to separate them.

Paratenetus fuscus
Diagnosis. This species differs from P. gibbipennis by the character states listed in the description.
Description. Same character states as P. gibbipennis except for the following: slanting setae on elytra less depressed, semierect, occasionally even suberect; erect setae numerous, in seven or eight rows; metaventrite slightly longer, length along midline subequal to slightly shorter than length of abdominal ventrite 2 along midline.
Specimens were collected in February (n=1), March (n=30), April (n=58), May (n=8), June (n=40), July (n=7), August (n=19), September (n=14), October (n=5) and November (n=4). ". The first two specimens correspond neither to our concept of P. punctatus nor to any other North American species we have seen. The specimens are in poor condition, with almost all the setae gone, but they appear to be conspecific. Although Spinola indicated that all three of his specimens came from the United States and were provided by "Mr.

Paratenetus punctatus
[John Eatton] LeConte," these two specimens may have originated from Mexico, Central America or South America. The third specimen, a small individual (3.2 mm), fits our concept of P. punctatus and is here selected as lectotype. The label "Lectotype Paratenetus punctatus Spinola des. Y. Bousquet 2012" has been attached to the specimen.
Diagnosis. Many specimens of P. punctatus can be separated from the other North American species of Paratenetus by their large size (3 mm or more). The vast majority of specimens of the other species are less than 3 mm long. Otherwise, the species can be separated from P. exutus in having the antennomere 8 subquadrate, the pronotum wider clearly anterior to the midlength, the punctation on the pronotum coarser, the slanting setae on the elytra slightly longer and more erect and the protibia of the male with a calcar along ventral surface. From P. texanus, this species is best separated in having the pronotum widest anterior to the midlength and the punctures on the pronotum coarser and denser, in part subcontiguous along the lateral half.
Description. Body dorsally uniformly pale to dark reddish brown in most specimens, with the pronotum and head slightly darker than elytra and legs in some specimens; antennal club darker than antennomeres 1-8; metaventrite not darker than first two abdominal ventrites in the vast majority of specimens, slightly darker in a few specimens. Antennomere 8 subquadrate. Pronotum with maximum width anterior of midlength (Fig. 3); punctures narrowly spaced, in part subcontiguous over lateral half. Elytra less convex than for P. gibbipennis and P. fuscus; slanting setae semierect in the vast majority of specimens, suberect in some specimens, erect setae few. Metaventrite long, length along midline longer than length of abdominal ventrite 2 along midline. Male protibia with calcar near middle along ventral surface; male mesotibia with very short, preapical spine, oriented perpendicularly or obliquely to long axis of tibia. Parameres with sides more or less parallel to very slightly convergent towards apex; apex more or less rounded (Fig. 8).
Length: 3.0-4.0 mm. Distribution. This species ranges from New Brunswick to southeastern Manitoba, south to eastern Texas, southern Mississippi, and southeastern Florida (Fig. 12).

Paratenetus exutus
Etymology. The specific name comes from the Latin participle exutus, -a, -um (deprived of) and alludes to the fact that the protibia of the male lacks the spinelike projection (calcar) found in the other American (north of Mexico) species.
Diagnosis. This species is best separated from P. punctatus and P. texanus in having the antennomere 8 transverse. The males are also easily recognized among the species treated here in having no calcar on the protibia and a relatively long apical spine, oriented more or less parallel to long axis of tibia, on the mesotibia.
Description. Body dorsally pale reddish brown in most specimens, with the pronotum and head usually slightly darker than elytra and legs; antennal club darker than antennomeres 1-8, particularly in males; metaventrite quite distinctly darker than first two abdominal ventrites in the vast majority of specimens, not or only slightly darker in a few specimens. Antennomere 8 transverse. Pronotum with maximum width at or very slightly anterior of midlength (Fig. 2); punctures narrowly spaced, in part subcontiguous over lateral half. Elytra less convex than for P. gibbipennis and P. fuscus; slanting setae subdepressed, erect setae few. Metaventrite long, length along midline longer than length of abdominal ventrite 2 along midline. Male protibia without calcar near middle along ventral surface; male mesotibia with relatively long, apical spine, oriented more or less parallel to long axis of tibia. Parameres with sides convergent towards apex; apex more or less truncate (Fig. 7).
Length: 2.5-3.0 mm. Detroit (USNM). Minnesota. Becker Co.: Itasca St. Pk. area (USNM). Etymology. The specific name derives from the name of the state of Texas where the species has been commonly collected.
Diagnosis. Members of this species can be distinguished from those of P. punctatus and P. exutus in having the punctures on the pronotum sparser, not subcontigous even on the lateral half. They can also be distinguished from most adults of P. punctatus by their smaller size and from most adults of P. exutus by the subquadrate antennomere 8 and metaventrite of same color as the first two abdominal ventrites.
Description. Body dorsally yellow to pale reddish brown, with the pronotum and head usually slightly darker than elytra and legs; antennal club slightly darker than antennomeres 1-8 in many specimens, often reddish brown to partially piceous, yellowish and as pale as legs in some specimens; metaventrite not darker than first two abdominal ventrites. Antennomere 8 subquadrate. Pronotum with maximum width at midlength (Fig. 4); punctures moderately dense, not subcontiguous even over lateral half. Elytra less convex than P. gibbipennis and P. fuscus; slanting setae subdepressed, erect setae short. Metaventrite long, length along midline longer than length of abdominal ventrite 2 along midline. Male protibia with calcar near middle along ventral surface; male mesotibia with short, preapical spine, wide at base and oriented perpendicularly to long axis of tibia. Parameres with sides distinctly convergent towards apex; apex markedly acute (Fig. 9).
Length: 2.7-3.3 mm. Distribution. This species is known from southeastern Florida, central Louisiana, and central and eastern Texas (Fig. 14). We have also seen specimens from the states of Chiapas, Nayarit and Tamaulipas in Mexico.
Records. We have seen 515 specimens, including the type material, from the following localities. United States of America. Florida. Dade Co.: Miami (FSC). Louisiana. Avoyelles Parish: Mansura (USNM). Cameron Parish: Holly Beach (LSAM, TAMU); nr. Oak Grove (TAMU). Texas. "60 mi SE Cotulla" (CNC). "15 mi SW Jct   This new species occurs in Mexico and nine species have been reported from that country. We have examined the type material of the six species described by Champion and housed in BMNH, i.e., Paratenetus constrictus, P. corticarioides, P. nigricornis, P. punctulatus, P. tibialis, and P. villosus, and none of them are conspecific with those of P. texanus. The three species not seen are P. tropicalis Motschulsky, P. koltzei Pic, and P. mexicanus Pic.