Revision and hypothetical phylogenetic analysis of the species of the New World genus Ataeniopsis (Coleoptera: Aphodiinae: Eupariini)

The New World genus Ataeniopsis Petrovitz, 1973 is revised. Fifteen species are recognized including three new species: Ataeniopsis carupanoi sp. n. from Venezuela, A.jaltipani sp. n. from Mexico and A. vinacoensis sp. n. from Argentina. Lectotype of A. haroldi (Steinheil, 1872) is designated, the name of type species A. notabilis Petrovitz, 1973 is reestablished, five species are given in a new combination. The taxa are diagnosed, keyed and illustrated, and biological information and distribution data summa­ rized following the species descriptions. A hypothetical phylogenetic analysis of Ataeniopsis based on cladistic analysis is presented.


INTRODUCTION
This revision of the genus Ataeniopsis Petrovitz, 1973, is one of a series of papers published on the Neotropical Eupariini (Stebnicka, 1999a(Stebnicka, , 1999b(Stebnicka, , 2000(Stebnicka, , 2001a(Stebnicka, , 2001b(Stebnicka, , 2001c(Stebnicka, , 2002)).The genus with the type species Ataen iopsis notabilis Petrovitz, 1973 from Brazil, consists of a species complex structurally distinct from other genera of the Eupariini.Woodruff (1973) and Cartwright (1974) placed the North American species in the genus Ataenius Harold, 1867. Chalumeau (1992) treated Ataeniopsis as a subgenus of Ataenius, however, a number of consistent characters appear to warrant generic status.The genus contains fifteen currently recognized species ranging in distribution from the United States to Argentina.Seven species are found in South America, four in Mesoamerica and in the Sonoran province, three species occur exclu sively in the southeastern United States and one species inhabits the West Indies.This study considerably extends the distribution of Ataeniopsis which is presented in the form of maps, with the exception of species inhabiting the United States of America cited by Cartwright (1974).A generic diagnosis is given below, followed by a cladistic analysis, keys and descriptions.Since the species of Ataeniopsis are not clearly differentiated, their descrip tions include mainly: exceptions to the usual conditions mentioned in the generic diagnosis, the state of characters varying widely in the genus and unique features.
Remarks.Ataeniopsis is most closely related to Ataenius but exhibits a number of apomorphies, some unique and some in combinations that universally apply to all species of the genus.The best characters combina tions that make members of Ataeniopsis easy to recognize are: the elongate and highly polished body, the abdomi nal sternites with very fine sutures, the pygidium convex, shiny with no scabrous sculpture, and all femora lack marginal lines.
The vast majority of specimens from Central and South America were collected using light traps, which provide some information on their flight period but very little on their microhabitat.

Phylogenetic hypothesis
On the basis of the characters judged to be derived the genus Ataeniopsis is regarded as a monophyletic group.Fifteen species of Ataeniopsis form ingroup and five spe cies belonging to Ataenius texanus-group the outgroup (synapomorphies: size and shape of the body, sculpture of the head).The 22 characters with 47 character states hypothesized to be primitive and derived, are defined and coded in Table 1.

Table 1
Matrix of taxa and character states used in the cladistic analysis.

Synapomorphies for the following main lineage of Ataeniopsis are:
Character 4 -clypeal wrinkles fine.This state occurs in most Ataeniopsis species except for the parallelus + rugopygus + pusillus node, which is not supported by synapomorphies.These three species have shagreened and usually roughly sculptured head like the outgroup taxa.Character 5 -vertex of head with scattered punc tures; synapomorphy for nodes formed by carupanoi + regulus + notabilis and jaltipani + armasi + haroldi + vinacoensis.Plesiotypically, the punctures on vertex are dense and occur in a regular band; this character state concerns the nodes formed by two Sonoran species duncani + parkeri and by three Nearctic species edistoi + figurator + saxatilis.Character 12 -elytral intervals smooth, impunctate.The apomorphic state that typifies most species of Ataeniopsis; synapomorphy with rever sals in vinacoensis and figurator, which have the elytral intervals with more or less visible punctures.Character 1 -body slightly arcuate, not quite parallel-sided.The apo morphic character state and synapomorphy for the node formed by edistoi + figurator + saxatilis, the three spe cies known exclusively from the southeastern United States.The remainder of the species in the ingroup and outgroup have a slender and parallel-sided body.The spe cies of Ataeniopsis share a number of character states that are probably plesiomorphic and the retention of which serves as a reflection of their relatedness.ally separated by 2-3 diameters but variable in density, middle of lateral area lacks punctures or only very fine punctures visible.Elytra relatively long, parallel, striae usually distinctly impressed with punctures slightly crenating inner margins of intervals; discal intervals flat, shiny, minute punctures scattered or lacking.Prosternum and inner side of profemur with long pale hair; metasternal midline shallow, sometimes disc of metasternum slightly concave; abdominal sternites dis tinctly fluted along sutures, surface slightly swollen at extreme sides.Profemur, meso-and metafemur fusiform, equal in length; metatibia slender, apex in some speci mens with very minute accessory spine; basal tarsomere of metatarsus equal in length to following three tarsomeres combined.Epipharynx as in Fig. 4.
Female.Body usually more elongate than in male; abdominal sternites 2-4 equal in length, sternite 5 half length of sternite 4.
Remarks.Ataeniopsis haroldi is one of the larger spe cies of the genus occurring south of the Amazon basin (Fig 12).It was considered as a widely distributed species, recorded by Chapin (1940) from Mexico, Cuba, Hispaniola, Puerto Rico and North America.Deloya (1994) followed Chapin in citing this misleading distribu tion without critical comment.As indicated on the labels, the specimens were collected to light traps in subtropical seasonal forest and in riverine forest and beaten from leaves.
Remarks.Variation in Ataeniopsis regulus is consider able, both within a locality and between widely separated localities.The populations from Ecuador differ from those of Venezuela, showing variation in size and in the punctuation of pronotum and may at first glance appear to be separate species.Such variation is observed among other species of the Neotropical Aphodiinae and it is undoubtedly due to the different environmental condi tions.A. regulus is one of the most common species of Ataeniopsis, closely related to A. carupanoi sp.n. and to A. notabilis (see Remarks under those species).All speci mens with collection data were taken at UV lights, on coastal area of lagoon and lakes, in coconut groves.Petrovitz, 1973 (Figs 8, 13) Ataeniopsis notabilis Petrovitz, 1973: 191.-Dellacasa 1988: 170 (catalogue).Ataenius (Ataeniopsis) notabilis : Chalumeau 1992: 196 (as synonym of A. regulus).Diagnostic characters.Length 3.2-3.8mm.Colour reddish black to black, legs usually lighter than bottom surface.Clypeal emargination shallow, denticles usually very fine, surface of head finely microreticulate, subo paque, frontal suture not indicated, clypeal wrinkles fine, punctures on vertex variable in density, usually irregu larly spaced.Pronotum slightly transverse, basal marginal line absent, basal edge finely crenate, crenations visible when pronotum viewed from behind; pronotal punctures variable, usually fine along anterior margin and larger towards base and sides, mixed fine and moderate, irregu larly spaced, fine and shallow at anterior angle and along lateral margin.Elytra parallel, about 2.5 times as long as pronotum, striae shallow with fine punctures; discal inter vals flat, shiny, surface punctures very minute.Pros ternum and inner side of profemur with short hair; metasternal midline shallowly impressed; abdominal sternites without fluting, almost smooth along sutures, sur face smooth, impunctate; pygidium with 3-4 fine punctures along median carina, apical lip convex.Mesoand metafemur equal in length, fusiform with fine scat tered punctures; metatibia slender, apex without acces sory spine; basal tarsomere of metatarsus a trifle longer than following three tarsomeres combined.
Remarks.Chalumeau (1992) synonymized Ataeniopsis notabilis with A. regulus.On the basis of a careful study of the external characters and the male genitalia, I am here reestablishing the name A. notabilis as the type spe cies of the genus.The species differs from regulus in having more distinct clypeal wrinkles, pronotum lacking a marginal line and abdominal sternites nearly smooth, without fluting.(Burmeister, 1877), comb.n. (Figs 2,9,13) Euparia pusilla Burmeister, 1877: 410.Ataenius pusillus : Schmidt 1908: 94;1922: 434;Dellacasa 1988: 280 (catalogue).Diagnostic characters.Length 3.5-3.8mm.Colour of body (Fig. 2) piceous, legs lighter Clypeal emargination moderate, edge on each side of emargination obtuse, even slightly rounded, lacking denticles; surface of head microreticulate, usually roughly wrinkled from anterior margin to vertex, occipital area with band of close punc tures.Pronotum slightly transverse, base distinctly mar gined by smooth line; pronotal punctures fine to moderate in size, on disc irregularly spaced becoming closer toward sides and lateral margin, separated by about one diameter.Elytra parallel, about 2.5 times as long as pronotum, striae distinctly impressed with fine punctures; discal intervals flat, shiny, surface punctures minute to fine, scattered, sometimes invisible.Prosternum and inner surface of pro femur with close whitish hair; mesosternum rugosely shagreened, metasternal midline shallow; abdominal sternites finely fluted along sutures, surface smooth, impunctate, slightly swollen at extreme sides, pygidium polished, con vex, in some specimens disc with trace of longitudinal carina and few minute punctures.Meso-and metafemur fusiform, with fine scattered punctures; metatibia slender, apex without accessory spine; basal tarsomere of meta tarsus a trifle longer than following three tarsomeres com bined.
Remarks.Ataeniopsis pusillus as now defined is one of the commonest members of the genus.The shape of clypeus and sculpture of the head as well as its southern distribution in South America (Fig. 13) easily distinguish this species.It seems to be closest to A. parallelus, but differs from that species by the characters given in the key.Specimens were collected in the area of suburban grassland, riverine forest and tropical transition forest, attracted to UV light traps in a great number of individu als.
Diagnostic characters.Length 2.8-3.0 mm.Body moderately shining, piceous black, legs reddish brown.Clypeal emargination shallow, denticles on each side of emargination fine but distinct; surface of head subopaque, finely, roughly sculptured, frontal suture marked by line, vertex with band of close punctures.Pronotum slightly transverse, basal marginal line fine with minute crenations; pronotal punctures mixed minute and fine, very close, everywhere distributed.Elytra parallel, about 2.5 times as long as pronotum, striae deep with slightly crenating punctures; discal intervals slightly convex, surface punctures very minute.Prosternum and inner surface of profemur with short hair; metasternal midline shallow; abdominal sternites finely fluted along sutures, surface slightly uneven at extreme sides; pygidium with 3-4 fine punctures along median carina, apical lip convex.Mesoand metafemur fusiform with fine scattered punctures, metafemur a trifle longer than mesofemur; metatibia slen der, apex without accessory spine; basal tarsomere of metatarsus longer than following three tarsomeres com bined.
Remarks.The small size and very close punctures on the pronotum easily distinguish A. parallelus from related species.No additional specimens were found in the mate rial examined and nothing is known about its biology.Description.Length 4.2-4.5 mm.Colour dark red to black, legs reddish brown.Head markedly gibbose at middle, clypeus with shallow emargination and sharp denticles on each side; surface of head shiny, finely wrin kled to vertex, frontal suture not indicated, punctures on vertex very fine, variable in density, usually irregularly spaced.Pronotum subquadrate, basal marginal line dis tinct, basal edge smooth; pronotal punctures mixed min ute, fine and moderate, usually fine along anterior margin then larger towards base and sides, irregularly spaced, shallow at anterior angle and vanishing at posterior angle.Elytra parallel, slender, about 3 times as long as pronotum, striae distinctly impressed with fine contiguous punctures slightly crenating inner margins of intervals; discal intervals slightly convex, shiny, surface punctures very minute scattered or lacking.Prosternum and inner surface of profemur with rather long hair; metasternal midline impressed; abdominal sternites with moderate fluting along sutures, slightly arcuate, surface smooth, impunctate; pygidium polished, slightly curved ventrally, apical lip convex.Meso-and metafemur equal in length, fusiform with fine scattered punctures; metatibia slender, apex without accessory spine; basal tarsomere of meta tarsus longer than following three tarsomeres combined.Epipharyngeal structures similar to those of A. haroldi (Fig. 4).
Description.Length 3.2-3.6mm.Body slender, red dish black, legs somewhat lighter.Clypeal emargination shallow, denticles on each side of emargination fine, sur face of head shiny, distinctly wrinkled from anterior margin to vertex, punctures on vertex fine, not close, irregularly spaced.Pronotum subquadrate, basal marginal line very fine, basal edge minutely crenate, crenations visible when pronotum viewed from behind; pronotal punctures moderate in size, slightly irregularly distributed, separated by 2-3 diameters, lacking along lat eral margin.Elytra narrow, parallel, about 2.5 times as long as pronotum, striae shallow with fine punctures; discal intervals flat, shiny, surface punctures invisible.Prosternum and inner side of profemur with rather long pale hair; metasternal midline shallowly impressed; abdominal sternites with inconspicuous fluting, almost smooth along sutures, surface smooth, impunctate; pygidium polished, apical lip convex.Meso-and metafemur fusiform with fine scattered punctures, metafemur a trifle longer than mesofemur; metatibia slen der, apex without accessory spine; basal tarsomere of metatarsus longer than following three tarsomeres com bined.Epipharyngeal structures similar to those of A. haroldi (Fig. 4).
Affinity.The species is most closely related to A. regu lus, but surface of the head is not so distinctly shagreened as in A. regulus, the pronotal lateral area is almost impunctate and the abdominal sternites are inconspicu ously fluted.
Ataeniopsisjaltipani sp.n. (Figs 12,18) Type material.Holotype male, Mexico, Veracruz, 3.5 km E Jaltipan, Rte.Description.Length 3.2-3.8mm.Body parallel, colour reddish black to black, legs usually lighter than bottom surface.Clypeus almost truncate anteriorly or emargination very shallow, denticles on each side of emargination acutely triangular, surface of head finely microreticulate, shining, frontal suture not indicated, clypeal wrinkles fine, punctures on vertex variable in density, not close, usually irregularly spaced.Pronotum slightly transverse, basal marginal line absent, basal edge almost smooth, pronotal punctures mixed minute and fine, the latter gen erally separated by more than one diameter, shallow at anterior angles, scattered on sides, minute punctures visible along lateral and anterior margin.Elytra parallel, about 3 times as long as pronotum, striae impressed with fine punctures; discal intervals flat, shiny, surface punc tures indistinct.Prosternum and inner surface of profemur with moderate to long hair; metasternal midline shallow; abdominal sternites arcuate without fluting, almost smooth along sutures, surface smooth, impunctate; pygidium polished, strongly shining, slightly curved ven trally, apical lip convex.Meso-and metafemur fusiform with fine scattered punctures; metafemur slightly longer than mesofemur; metatibia slender, apex without acces sory spine; basal tarsomere of metatarsus subequal in length to following three tarsomeres combined.Epipha ryngeal structures similar to those of A. haroldi (Fig. 4).
Female.Abdominal sternites 3 and 4 equal in length; each slightly longer than sternite 5.
Affinity.Ataeniopsis jaltipani sp.n. is closely related to A. duncani, but may be easily distinguished by the lack of basal marginal line of the pronotum and nearly smooth abdominal sutures.The present state of knowledge indi cates that these species are allopatrically distributed (Fig. 12).
Diagnostic characters.Length 3.3-3.8mm.Colour reddish black to black, legs somewhat lighter.Clypeal emargination shallow, denticles on each side of emargina tion fine but distinct; surface of head shiny, finely wrin kled from anterior margin to vertex, punctures on vertex variable in density, usually irregularly spaced.Pronotum slightly transverse with basal marginal line, basal edge smooth; surface punctures moderate in size, usually finer along anterior margin and larger towards base and sides, irregularly spaced, fine and shallow at anterior angle and missing at posterior angle.Elytra parallel, about 3 times as long as pronotum, striae impressed with contiguous, crenating punctures; discal intervals slightly convex, sur face punctures very minute.Prosternum and inner side of profemur with rather long hair; metasternal midline weakly marked; abdominal sternites 2-3 without fluting, sternites 4-5 finely fluted along sutures, surface smooth, impunctate; pygidium polished, apical lip convex.Mesoand metafemur equal in length, both with fine scattered punctures; metatibia slender, apex without accessory spine; basal tarsomere of metatarsus longer than fol lowing three tarsomeres combined.
Remarks.The West Indian Ataeniopsis armasi is inter mediate between the Mesoamerican species A. duncani and A. jaltipani sp.n.It differs from the former species by its less heavily punctate pronotum, from the latter by partially fluted abdominal sternites and presence of a pro notal basal margin.Chalumeau (1982) recorded this spe cies from Puerto Rico, Dominican Rep., Haiti and Guadeloupe.The specimens were collected to black light traps.
In both sexes, abdominal sternite 3 longer than sternites 4 and 5; sternites 4, 5 equal in length.In male, pygidium longer and more convex than in female, genitalia as in Fig. 17.
Remarks.Ataeniopsis rugopygus may be easily recog nized by the characters given in the key.Some specimens show uneven pygidial surface, and some have the pygidium with very little other sculpture.The species occurs in the Sonoran province (Fig. 12), recorded by Cartwright (1974, Fig. 4) from Arizona, Colorado, Iowa, New Mexico and Texas.In Mexico found for the first time.
In both sexes, abdominal sternite 3 longer than sternites 4 and 5; sternite 5 half length of sternite 4. In male, disc of pygidium longer and more convex than in female, genitalia as in Fig. 20.
Remarks.Ataeniopsis figurator was synonymized with A. haroldi by Chapin (1940) who cited it from Argentina, Mexico, West Indies and North America.This species, widely distributed in the middle and eastern United States (see Cartwright 1974, Fig. 5), is apparently restricted to that area and does not occur in Mexico.The distribution data given by Deloya (1994) is that of at least three other species.
Remarks.Ataenius saxatilis is most close to A. figu rator but it is more robust than A. figurator and than all other species in the genus, and differs in a general shape.Apart from the type series, no additional specimens were found in the material examined.This species was reported by Cartwright (1974, Fig. 4) from Georgia and North Carolina, specimens were collected from the sand and soil around weathered areas of granite and gneissic rocks.(Cartwright, 1974), comb.n.Ataenius edistoi Cartwright, 1974: 17-18, Fig. 2. -Dellacasa 1988: 343 (catalogue).Diagnostic characters of female.Length 3.9 mm.Elongate-oblong, shiny black with anterior margin of clypeus, anterior margin of pronotum and legs reddish.Clypeal emargination wide and shallow, edge on each side rounded, with no trace of denticles, surface of head wrinkled over anterior half, upper half closely finely punctate, vertex with narrow band of fine punctures.Pro notum subquadrate, basal marginal line distinct; surface with evenly distributed, close, mixed very fine and mod erate punctures generally separated by about one diameter, more numerous at anterior angle.Elytra not quite parallel, about 2.5 times as long as pronotum, humeri finely dentate; striae moderately crenate-punctate, discal intervals flat, shiny, surface punctures very minute.Prosternum and inner surface of profemur with moder ately long hair; metasternal midline long, not deeply impressed, disc finely punctate; abdominal sternites finely fluted along sutures, surface punctures fine, scattered, sternites 2-4 equal in length, sternite 5 about three quar ters length of sternite 4; pygidium with fine, rather close punctures along median carina.Meso-and metafemur equal in length with fine scattered punctures; apex of metatibia expanded without accessory spine; basal tarso mere of metatarsus a trifle longer than following three tar someres combined.

Ataeniopsis edistoi
Remarks.Ataeniopsis edistoi is closely related to A. figurator and A. saxatilis and similar in general appear ance, being slightly broader in shape than figurator and not quite as broad as saxatilis.However, the lack of male specimens in the material examined (except a single available, badly damaged individual) does not allow a comparison of male genitalia.A. edistoi is hitherto known from a small number of specimens collected exclusively in South Carolina (Cartwright 1974, Fig. 2) under surface litter on sandy ground along the roadsides.

Fig. 12 .
Fig. 12. Known distribution of Ataeniopsis in Central and South America.
Fig. 13.Known distribution of Ataeniopsis in Central and South America.