A comprehensive guide to the Argentinian case-bearer beetle fauna (Coleoptera, Chrysomelidae, Camptosomata).

Knowledge of Argentinian Camptosomata has largely remained static for the last 60 years since the last publication by Francisco de Asis Monrós in the 1950's. One hundred and ninety Camptosomata species (182 Cryptocephalinae and 8 Lamprosomatinae) in 31 genera are recorded herein from Argentina. Illustrated diagnostic keys to the subfamilies, tribes, subtribes and genera of Argentinian Camptosomata, plus species checklists and illustrations for all genera of camptosomatan beetles cited for each political region of Argentina are provided. General notes on the taxonomy and distribution, as well as basic statistics, are also included. This study provides basic information about the Camptosomata fauna in Argentina that will facilitate in the accurate generic-level identification of this group and aid subsequent taxonomic revisions, and phylogenetic, ecological, and biogeographic studies. This information will also facilitate faunistic comparisons between neighboring countries. Two nomenclatural acts are proposed: Temnodachrys (Temnodachrys) argentina (Guérin, 1952), comb. n., and Metallactus bivitticollis (Jacoby, 1907), comb. n. The following are new records for Argentina: Stegnocephala xanthopyga (Suffrian, 1863) and Lamprosoma azureum Germar, 1824. Currently, the most diverse camptosomate tribe in Argentina is Clytrini, with almost twice the number of species of Cryptocephalini. New records for Argentina are predicted.

Introduction

In Argentina there are 956 recorded species of (excluding ) in 258 genera (Cabrera and Roig-Juñent 1998a). Only two subfamilies of leaf beetles are not represented in Argentina: Kirby and LeConte and Horn. Most subtropical species are distributed in the Amazonian and Chacoan domain (Cabrera and Willink 1973). However, current knowledge of Argentinian is incomplete. The present paper is the first one in a planned series on the Argentine chrysomelid fauna.

The chrysomelid subfamilies and are collectively known as “Camptosomata” or “case-bearers,” due to the peculiar habit of having their eggs, larvae, and pupae living in a fecal protective case (Brown and Funk 2005; Chaboo et al. 2008; Erber 1988, Jolivet and Hawkeswood 1995). Adults of case-bearing chrysomelids feed on foliage of a variety of eudicots (Erber 1988), but their larvae often show departures from strict phytophagy. The larvae of some and live in ant nests, where they feed on other items such as ant droppings and pellets, detritus, leaf litter and even dead insects collected by the ants (Agrain et al. 2015, and references therein). The larvae of camptosomates can be easily recognized by the behavior of carrying a portable case and the J-shaped body morphology.

includes four tribes (Chamorro and Konstantinov 2011): (1 genus), (2 genera), (1 genus), and (10 genera) (Seeno and Wilcox 1982), totalling 190 described species (Reid 2016). Reid (2016) and Chamorro (2014a), concur on a world estimate of 250 species. In Argentina, the only genus represented is Kirby. includes ~5300 species, independently calculated by Chamorro, (2014b) and Reid (2016) that are classified into three tribes: , , and (until recently treated under the name ) as originally proposed by Reid (1995, 2000). Members of the subfamily are distributed worldwide, but many tribes have distinct distributions (Erber 1988). Species are phytophagous in the adult stage, primarily leaf and flower feeders. All three tribes of this subfamily have representative genera in Argentina. The main goal of this contribution is to provide an updated systematic framework for Argentinian Camptosomata, treating all of its genera in order to better measure our current knowledge of these groups. This work includes the compilation of former fragmentary literature on the subject.

Type material of Argentinian Camptosomata

Most of the type specimens of Argentinian Camptosomata are deposited in European institutions: The Natural History Museum, London, United Kingdom (BMNH), Hungarian Natural History Museum, Budapest, Hungary (HNHM), Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium (KBIN), Museo Regionale di Scienze Naturali di Torino (MRSN), Museum für Naturkunde der Humbolt Universitat, Berlin, Germany (ZMHB), and National Museum, Prague, Czech Republic (NMPC). There are also type specimens in institutions in the United States: Museum of Comparative Zoology Collection, Harvard University, Boston, USA (MCZ), and the National Museum of Natural History, Smithsonian Institution, Washington D.C., USA (USNM). Yet, some type specimens are deposited in Museo de La Plata, La Plata, Argentina (MLPA) (see Cabrera and Roig-Juñent 1998b), Instituto y Fundación Miguel Lillo, Tucumán, Argentina (IMLA) (Aranda et al. 2016), and Museo Argentino de Ciencias Naturales ‘Bernardino Rivadavia’, Buenos Aires, Argentina (MACN) (Bachmann and Cabrera 2010). Two of the most prominent workers on Argentinian Camptosomata were Francisco de Asis Monrós, whose collection was donated to the Smithsonian Institution (USNM) (Staines 1995), and Manuel Viana, whose collection is now housed in Tucumán and Salta Provinces in Argentina and in Chile. More recently, a few type specimens have been deposited in in the Instituto Argentino de Investigaciones de las Zonas Áridas, Mendoza, Argentina (IADIZA) by Agrain (2013, 2014).

Methods

We studied all catalogs and specialized literature dealing with the genera treated in this contribution. Nomenclature follows previous authors, especially those who made extensive revisions of this group, such as Andrew Moldenke, Francisco Monrós, Jacintho Guérin, and Martin Jacoby. Characters used for identification keys are those used by: Agrain and Roig-Juñent (2011), Chamorro-Lacayo and Konstantinov (2009), Guérin (1943), Karren (1972), Lacordaire and Chapuis (1854), Moldenke (1970, 1981), Monrós (1949a, 1953a), and Riley et al. (2002). An identification key to the subfamilies, tribes, subtribes, genera, and subgenera of Argentinian Camptosomata was made by compiling and modifying previous publications as indicated in Table 1. Some couplets in our key, derived from keys provided by earlier authors, are based on extreme representatives of a rather continuous spectrum. The latter is due to the fact that many genera, and especially subgenera, require modern revision. Our key is built for the identification of taxa on the territory of Argentina but is useful for the South American continent. The characters given for some widely distributed genera (e.g., Geoffroy, Chevrolat) may not apply to species outside Argentina. Images of dorsal and lateral habiti were taken by different authors as indicated in superscript values: (1) F. Agrain, (2) L. Chamorro, (3) C. Gorretta, N. Cabrera, and (4) D. Sassi, and edited by F. Agrain.

Table 1.

Main sources of information used for the identification key.

Group	Citation
Genera of Fulcidacini	Chamorro-Lacayo and Konstantinov (2009), Karren (1972), Lacordaire and Chapuis (1854).
Genera and subgenera of Clytrini	Agrain and Roig-Juñent (2011), Lacordaire and Chapuis (1854), Moldenke (1981), Monrós (1953a).
Subtribes of Cryptocephalini	Lacordaire and Chapuis (1854), Riley et al. (2002).
Genera of Cryptocephalina	Lacordaire and Chapuis (1854), Monrós (1949a), Watts (2005).
Genera of Pachybrachina	Chamorro (2013); Lacordaire and Chapuis (1854).

We conducted an exhaustive search of all publications citing Argentinian camptosomates. Here we present a checklist of all currently known camptosomate species from Argentina, their distribution, host plant preferences, juvenile data where available, and known predators. Junior synonyms are provided for each species when applicable. The 24 provinces in Argentina (Fig. 1A) are abbreviated as follows: Buenos Aires (BAS), Catamarca (CAT), Chaco (CHA), Chubut (CHT), Ciudad Autónoma de Buenos Aires (CAB), Córdoba (COR), Corrientes (CTS), Entre Ríos (ERS), Formosa (FOR), Jujuy (JUY), La Pampa (LPA), La Rioja (LRA), Mendoza (MZA), Misiones (MNS), Neuquén (NQN), Río Negro (RNO), Salta (SAL), San Juan (SJN), San Luis (SLS), Santa Cruz (SCZ), Santa Fe (SFE), Santiago del Estero (SEO), Tierra del Fuego (TFO), Tucumán (TUC). The source map of Andean and Neotropical regions of Argentina was obtained from Löwenberg-Neto (2014).

Figure 1.

Distribution of Camptosomata tribes. A Map showing Argentinian administrative divisions with abbreviation as used in text. Andean and Neotropical regions as indicated in color reference B In black, administrative divisions containing species C In black, administrative provinces containing species D In black, administrative divisions containing species E In black, administrative divisions containing species.

Terminology

Terminology follows previous authors as indicated in table 1. The term “egg anal pit” refers to a medioventral excavation on terminal abdominal ventrite, mostly present in the females (Chaboo 2007, and references therein). The term “rectal apparatus” refers to a specialized region of the rectum of females, whose sclerites and muscles are used during oviposition (De Monte 1957, Erber 1988, Brown and Funk 2005, Schöller 2008). This structure is unique to the (Reid 1995).

Results

Keys to the subfamilies, tribes, subtribes, genera, and subgenera of Argentinian Camptosomata

Note: Some of the characters in this key are valid for Neotropical species only

Diagnostic characters plate 1. A Body dorsally highly convex and ventrally flattened (semicircular in cross-section (drawn after Monrós 1956) B Distal margin of last ventrite thick (drawn after Chamorro and Konstantinov 2011) C Last ventrite not excised in shape of arc D Body cylindrical, not flattened ventrally (drawn after Monrós 1953a) E body surface usually tuberculate (drawn after Monrós 1951) F procoxae prominent and contiguous G procoxae not prominent and separated H prosternal process more than 3/4 as long as intercoxal prosternal process (after Chamorro-Lacayo and Konstantinov (2009) I intercoxal prosternal process gradually constricted at about 2/3 of its length (after Chamorro-Lacayo and Konstantinov (2009).

Diagnostic characters plate 2. A Tarsal claws bifid or appendiculate B Frons with deep transverse sulcus. C Head forming straight angle with respect to pronotum D Head strongly directed downward, forming 45º angle with respect to pronotum E Posterior margin of pronotum broadly expanded, forming distinct scutellar lobe with angular corners (drawn after Monrós 1953a) F Pronotum margined at base, not crenulate G Pronotum not margined at base, crenulate.

Gyllenhal, 1813

Adults: Body cylindrical, or rarely as long as wide; in dorsal view parallel-sided with prothorax mostly as wide as combined elytral bases; rarely body rounded; multicolored and patterned, particularly , black with red humeri commonly in , brown, black, straw-yellow and some with velvet spots in , glabrous to pubescent, particularly . Head retracted into prothorax up to frons or almost completely, with compound eyes completely to barely visible from above. Compound eyes entire, level to strongly protuberant; canthus weak to deep. Antennae 11-segmented, longer than pronotum and filiform in (sometimes antennomeres distally dilated and flattened), shorter than pronotum and dentate in , clavate in . Pronotum about 0.75–1.0 times as long as wide, widest basally; sides slightly rounded or sinuate; base slightly narrower or as wide as combined elytral bases. Prosternum in front of coxae usually narrow and shorter than shortest diameter of a single coxal cavity, flat to moderately convex, sometimes produced to conceal mouthparts. Prosternal process complete, usually parallel-sided; notosternal sutures distinct. Procoxae not projecting below prosternum, without concealed lateral extensions; trochantins exposed within coxal cavity. Stridulatory device present on concealed part of mesoscutellum. Tarsi 5-5-5 in both sexes; penultimate tarsomere reduced and antepenultimate bilobed, all usually wider in males; tarsomere III densely clothed beneath with adhesive microtrichia; pretarsal claws simple to deeply bifid. Abdomen with five free ventrites and six tergites. Ventrite I more than twice as long as II, usually longer than ventrites II-IV combined, without postcoxal lines; intercoxal process narrowly rounded to almost truncate. Functional spiracles present on tergites I-VI. Tergite VI forming strongly pigmented pygidium, always exposed; anterior edge of sternite VIII in male without median strut. Ventrite V (=sternite VII) in females with variably-sized apical fovea. Males with segment IX membranous and spiculum gastrale Y-shaped. Aedeagus of cucujiform type; tegmen Y-shaped; struts (remnants of tergite IX) either present or absent; penis flattened to rounded, slightly to strongly curved apically; apically and/or laterally usually with tufts of setae. Sternite VIII in female lacking spiculum ventrale. Ovipositor short, rigid and oval PageBreakwith distinct proctigeral, paraproctal, and coxital baculi; paraprocts deltoid, slightly shorter than undivided coxites, sclerotized or less pigmented proximally, flattened, digitate lobes of variable form, apically setose; styli absent. Spermatheca strongly to moderately sclerotized, variably shaped, usually J-, C-, or S-shaped. Rectal sclerites (rectal apparatus) present in female.

Larvae: J-shaped, generally protected by a case. According to Reid (1990, 1995), Agrain and Marvaldi (2009) and Chamorro (2014b), the following features characterize the subfamily in the broad sense (i.e., including , and ), and are probably synapomorphies of cryptocephalines, by outgroup comparison with other chrysomeloids and weevils (Reid 1995, 2000): body J-shaped; frons, clypeus, and labrum fused; six stemmata, clustered 4 + 2; spiracles uniforous with reticulate peritreme; egg bursters on TII and TIII and associated with a long and a short seta. show a number of larval features in common with the cryptocephalines, being the body J-shaped among the most obvious (and related with their habit of carrying a case), as well as the fusion of frons, clypeus and labrum. Yet, unlike the cryptocephaline larvae, those of have bicameral spiracles with peritreme simple, and five stemmata grouped 2 + 3. The maxillary palp 3-segmented plus the palpiger, as present in both groups, is likely a plesiomorphy, and although both subfamilies have egg-bursters confined to the meso- and metathorax, those in lack the short ventral seta (Agrain and Marvaldi 2009, and references therein).

Tribe Kirby, 1837

Subtribe Lacordaire, 1848

Diagnosis. Same as for genus, see below.

Lacordaire, 1848

Fig. 4

Figure 4.

Lacordaire (1), left: habitus (dorsal view), right: habitus (lateral view).

Type species.

Lacordaire, 1848. By monotypy.

Diagnosis.

This genus is easily recognized by the presence of strong parallel carinae on the elytra; other diagnostic characters include: frons very narrow; scutellum slanting posterodorsally, distinctly protruding from the plane of elytra; intercoxal prosternal process nearly absent between fore coxae; epipleural margin broadly angled, tip rounded; elytra without pubescence; frons with medial pit, densely and coarsely punctate throughout; and pygidium strongly convex.

Distribution.

Only two species from Brazil, one present in Argentina, likely to also occur in Paraguay.

Remarks.

Agrain and Roig-Juñent (2011), found eight autapomorphies for the genus, among them elytra with strongly marked longitudinal striae constitutes an exclusive synapomorphy to the genus.

Argentinian species checklist.

1. Lacordaire, 1848 (FOR, MNS, SEO).

Subtribe Chapuis, 1874

Chevrolat, 1836

Chevrolat in

Gistel, 1848: 123.

Note.

Moldenke (1981), divided into six subgenera based on morphological features, such as the general shape of the body, pronotal margin, frons and antennomere morphology: Chevrolat and Moldenke from America north of Mexico; Moldenke, Moldenke, and Moldenke from the Neotropical region. Only the monotypic is represented in Argentina.

Moldenke, 1981

Fig. 5

Figure 5.

Guérin (1), left: habitus (dorsal view), right: habitus (lateral view).

Guérin, 1945. By monotypy.

This subgenus can be reliably diagnosed by the shape of anterior margin of pronotum, which is arcuate, and covers the entire head in dorsal view. Also, body shape is elongate (length 3x width), cylindrical and flat; frons is flat; lateral margin of prothorax not widely explanate.

Brazil and Argentina.

Moldenke (1981), mentioned the size of this species to be greater than 10 mm, but average size is smaller than 10 mm.

1. Guérin, 1945 (BAS, COR, ERS, JUY, LRA, RNO, SAL, SFE, TUC). Host plant: : sp. (Monrós 1953a).

Monrós, 1944

Fig. 6

Figure 6.

Monrós (2), left: habitus (dorsal view), right: habitus (lateral view).

Monrós (1944: 148).

This genus exhibits a particular combination of three characters unique among : adult body shape strongly cylindrically elongate; elytra not fully covering pygidium; and inconspicuous elytral punctations, thus superficially resembling a clerid.

This monotypic genus is limited to Paraguay and north and central Argentina.

According to Monrós (1944) this species has always been collected in extremely xeric places.

Monrós, 1944 (CAT, COR, MZA, SEO, SLS). Host plant: : (Quillo); : (Monrós, 1953a).

Erichson, 1847

Fig. 7

Figure 7.

(Erichson) (1), left: habitus (dorsal view), right: habitus (lateral view).

(Erichson, 1834), designated by Monrós (1953b: 48).

This genus resembles , but is distinguished by the epipleural lobe not well developed and from by the sides of the prothorax curved (strongly convergent towards the head); frons with deep medial pit; and distinct elytral pattern with transverse black bands on reddish base color.

Austral region of Chile and Argentina.

Monrós (1953b) separated the genera: , , and , formerly considered within . Currently monotypic.

(Erichson, 1834) (CHU, NQN, RNO). Host plant: : sp. (Monrós, 1953a) and (Roig-Juñent, 2004).

= Erichson, 1834.

= Lacordaire, 1848.

Lacordaire, 1848

Fig. 8

Figure 8.

Monrós (2), left: habitus (dorsal view), right: habitus (lateral view).

Lacordaire, 1848 designated by Monrós (1953a: 143).

Small body size (less than 7 mm). This genus is very similar to , from which it can be easily separated by the extraordinary development of the eyes, which are protruding and reaching the lateral margins of the head. Also, antennae with antennomere III large, conical; pronotum transverse; elytra without callus.

Brazil, Ecuador, Bolivia, Paraguay, and Argentina.

Monrós, 1953 (MNS).

Baly, 1877a (FOR, MNS).

= Baly, 1877a.

= Guérin, 1943.

Gistel, 1848

Fig. 9

Figure 9.

(Sahlberg) (1), left: habitus (dorsal view), right: habitus (lateral view).

Chevrolat, 1836

Lacordaire, 1848

Monrós, 1953a.

Sahlberg, 1823. By monotypy.

body elongate, brilliant metallic, uniform green/blue coloration; and bifid tarsal claws. Also, antennae serrated from IV antennomere; pronotal margin slightly marginate; scutellum long, triangular, with round apex; legs long, with long tarsi, last tarsomere surpassing ½ the lobes of tarsomere III.

Southern Brazil, Argentina and Paraguay. Only two subspecies of this genus have been cited in Argentina.

1a. (Sahlberg, 1823) (MNS).

= Sahlberg, 1823.

= Lacordaire, 1848.

1b. (Jacoby, 1897) (CHA, COR, JUY, MNS, SEO, TUC). Host plant: : , Argentina / Monrós, (1953a).

= Jacoby, 1897.

Moldenke, 1981

Fig. 10

Figure 10.

(Lacordaire) (1), left: habitus (dorsal view), right: habitus (lateral view).

This genus was created by Andrew Moldenke (1981) to include seven species previously included in . Moldenke (1981) divided it into two subgenera as follows:

Moldenke, 1981

Moldenke, 1981, by original designation.

According to Moldenke (1981), this group can be separated from by the following combination of characters: subrectangular body shape, with PageBreaksides subparallel; anterior margin of pronotum never concealing entire head in dorsal view; legs without longitudinal carinae. Additional characters that may help distinguish this genus are: frons without strong sexual dimorphism; antennomere IV clavate, smaller than V; hind pronotal angles sinuate, perpendicular.

Argentina, Brazil, Colombia, and Uruguay.

(Harold, 1875) (CHA, COR, CTS, LRA, MNS, MZA, SEO, SFE). Host plant: : (Mol.) Mol. (Viana and Williner 1972).

= Clavareau, 1913.

Monrós, 1953a (CHA, COR, FOR, JUY, MNS, SAL).

(Lacordaire, 1848) (CHA, COR, CTS, ERS, FOR, LRA, MNS, MZA, SAL, SEO, SFE, SJN, SLS, TUC). Host plant: : sp. (Monrós, 1953a), (Ward et. al., 1977) and (Roig-Juñent, 2004); (Viana and Williner, 1974); sp. (Ward et al., 1977).

(Monrós, 1953a) (CHA).

Moldenke, 1981 (JUY, SAL, FOR, CHA, MNS, SEO, TUC, CTS, COR, CAT, SFE, BAS, MZA). Host plant: : flowers (Roig-Juñent, 2004).

= Monrós, 1953a (nec Lacordaire, 1848).

Moldenke, 1981

Harold, 1875, designated subsequently by Moldenke (1981: 111).

Anterior margin of pronotum not explanate; male head very conspicuous; frons with very strong sexual dimorphism, male mandibles prominent; antennomere IV much smaller than V; hind pronotal angles obtuse.

This subgenus is endemic to Argentina.

(Monrós, 1953a) (CAT, CHA, COR, CTS, FOR, MNS, SEO, SFE, SLS). Host plant: : sp. and (Ward et al., 1977), (Viana and Williner, 1974).

(Monrós, 1953a) (COR, CTS, SFE, SLS).

(Harold, 1875) (BAS, CAT, CHA, COR, CTS, ERS, FOR, JUY, MNS, MZA, SAL, SEO, SFE, TUC). Host plants: : , and (Monrós, 1953a); (Aravena, 1940; 1974).

Lacordaire, 1848

Fig. 11

Figure 11.

Monrós (2), left: habitus (dorsal view), right: habitus (lateral view).

Lacordaire designated by Monrós (1953a: 150).

The most particular character to diagnose this genus is the shape of the head, which is anteriorly prolonged, strongly tapering and triangular. Other diagnostic characters are: elongate body, shiny and glabrous; eyes moderately salient; pronotal base lobate; scutellum long and,triangular.

Two species from Brazil, one of these with a subspecies in Northeastern Argentina which was separated from the typical form by Monrós (1953a) on the basis of its smaller size and distinct punctuation.

Monrós, 1953a (MNS, SFE).

Lacordaire, 1848

Fig. 12

Figure 12.

Lacordaire (1), left: habitus (dorsal view), right: habitus (lateral view).

Lacordaire, designated by Monrós (1953a): 215.

Sexually dimorphic, with males having enlarged forelegs; head completely concealed within prothorax; mesosternum broad; tarsal segments very transverse, tarsomere III cleft 1/2 length to receive IV; tarsomere IV notoriously short and PageBreakthick; frons with prominent transverse sulcation; lateral margin of pronotum broadly explanate, hind angles broadly rounded. This genus is closely related to , from which it can be separated by the conspicuous development of the legs, since the latter is much larger in males and females compared to .

Seven species and four subspecies are known from Argentina, another 10 species are known from Meridional America (southern part of South America).

() has been cited as a hostplant for sp. in Argentina (Monrós 1953a).

Lacordaire, 1848 (COR).

Lacordaire, 1848 (MNS).

3a. Harold, 1875 (CHA, COR, FOR, JUY, LRA, MNS, SAL, SEO, SFE, TUC).

= Harold, 1875.

3b. Lacordaire, 1848 (CHA, CTS, FOR, MNS, SFE). Host plant: : (Monrós, 1953a).

3c. Lacordaire, 1848 (JUY, SAL, SEO).

= Lacordaire, 1848.

4a. Monrós, 1953a (SFE).

4b. (Harold, 1875) (CAT, CHA, FOR, JUY, LRA, SAL, SEO, SLS). Host plant: : (Viana & Williner, 1974).

= Harold, 1875.

4c. Monrós, 1953a (JUY, SAL, TUC).

5. Monrós, 1953a (SEO).

6. Moldenke, 1981 (JUY). [Moldenke (1981), mistakenly cited this species from (Jujuy: Bolivia), it belongs to Jujuy: Argentina.].

Lacordaire, 1848

Moldenke (1981) erected to include North American species. The other four species of are included in the nominotypic subgenus and distributed in Central and South America, with only one species described from Argentina.

Lacordaire, 1848

Fig. 13

Figure 13.

Monrós (2), left: habitus (dorsal view), right: habitus (lateral view).

Lacordaire, 1848, designated by Monrós (1953a: 257).

This group can be easily distinguished from all other members of this subtribe by the very large and generally pointed epipleurae. South American species exhibit a black dorsal coloration always with metallic bronze reflections.

Usually foundin Central and South America.

Monrós, 1953a (BAS, CHA, COR, ERS, FOR, JUY, LPA, LRA, MZA, SAL, TUC).

Monrós, 1953

Fig. 14

Figure 14.

(Lacordaire) (1), left: habitus (dorsal view), right: habitus (lateral view).

Lacordaire, 1848 (part)

Monrós (1953b) divided this diverse genus (more than 60 species) into two subgenera based on the presence of a deep transverse sulcus in the interocular region.

Monrós, 1953

(Lacordaire, 1848), designated by Monrós (1953b: 49).

Frons without deep transverse sulcus; body minute, drop-like shaped with sides subparallel.

This subgenus has over 60 species and is distributed from Mexico to Argentina and Chile, 22 species have been cited for Argentina.

1a. (Lacordaire, 1848) (BAS, CHA, CTS, ERS, MNS, SEO, SFE, TUC). Host plant: : sp. (Roses) (Monrós 1953a).

1b. Monrós, 1953a (BAS, CTS, FOR, MNS, SFE).

(Lacordaire, 1848) (BAS, COR, MNS, MZA, SEO, SFE). Host plant: : sp. (Monrós, 1953a).

Monrós, 1953a (SEO).

Monrós, 1953a (NQN).

Monrós, 1953a (SFE).

Monrós, 1953a (JUY, SAL).

(Lacordaire, 1848) (JUY, MNS).

(Guérin, 1943) (FOR, MZA).

(Harold, 1875) (COR, MZA, SEO).

= Clavareau, 1913.

Monrós, 1953a (TUC).

Monrós, 1953a (COR, MNS).

(Monrós, 1951b) (MNS).

Monrós, 1953a (COR, MZA, SLS). Host plant: : (Ruiz & Pavon ex Hook.) Harms; (Roig-Juñent 2004).

Monrós, 1953a (JUY).

Monrós, 1953a (CHA, COR, SEO).

Monrós, 1953a (MNS).

(Lacordaire, 1848) (COR, JUY, SEO, TUC).

(Lacordaire, 1848) (CAT, SAL). Host plant: (Monrós 1947).

Monrós, 1953a (MNS).

Monrós, 1953a (CHA, COR, FOR, MNS, MZA, SAL, SEO, SFE, TUC). Host plant: : sp. (Monrós, 1953a).

Monrós, 1953a (JUY).

Monrós, 1953a (FOR, SEO, LRA, COR, SLS). Host plant: : Cav. (Quillo) (Monrós, 1953a).

Monrós, 1953

(Lacordaire, 1848), designated by Monrós (1953b: 49).

Frons with deep transverse sulcus; body shape subrectangular.

Seven species, from northern Brazil (one species) to central Argentina (six species).

The characters mentioned by Guérin (1952) in the original description of , are sufficient to transfer this species to this subgenus. The author mentioned the presence of a deep transverse sulcus in interocular region. Secondly, sides of the prothorax are much less convergent that expected and pronotal and elytral coloration pattern does not fit . In addition, distribution of is limited to Chile and southern Argentina, while Jujuy is the northernmost province. Finally, Guérin indicated the similarity with (Lacordaire), and Harold [= (Lacordaire)].

(Guérin, 1952), comb n. (JUY).

= Guérin, 1952.

(Lacordaire, 1848) (BAS, CHA, COR, CTS, ERS, FOR, JUY, LPA, MNS, MZA, NQN, SAL, SEO, SFE, TUC). Host plant: : , , and (Monrós 1953a); (Aravena 1974); . (Monrós 1953a).

= Harold, 1875.

= Lacordaire, 1848.

(Lacordaire, 1848) (BAS, COR, ERS, LPA, MNS, SFE). Host plant: : (Monrós 1953a).

Monrós, 1953a (ERS).

Moldenke, 1981 (CAT). Host plant: : (Cavanilles ex Lagasca) D.C., (Molina) Stuntz emend.; Griseb, (Gillies ex Hook. & Arn.) Benth.

Monrós, 1953a (JUY, SAL, CHA, LRA).

Monrós, 1953a (JUY, SAL).

(Lacordaire, 1848) (JUY, SAL, FOR, MNS, TUC, SEO, CAT, COR, LRA, CTS, SFE, SLS, BAS). Host plants: : (Mol.) Mol. (Monrós 1953a); Benth and (Griseb.) (Jolivet, 1978); (Cav.) Argentina / (Monrós 1953a); Anadenanthera colubrina var (Vell. Conc.) Brenan Argentina / (Jolivet 1978; Hayward 1944).

Lacordaire, 1848

Fig. 15

Figure 15.

Lacordaire (3), left: habitus (dorsal view), right: habitus (lateral view).

Moldenke (1981) divided this genus into five subgenera (including the nominotypic one). Two of these subgenera are present in Argentina, plus two species regarded by Moldenke (1981) as incertae sedis.

Moldenke, 1981

Lacordaire, 1848, designated by Moldenke (1981: 113).

Posterior margin of pronotum broadly expanded, forming distinct scutellar lobe usually bounded by acute angles; scutellum posteriad humeral callus, scutellum nearly as long as length of posterior lobe of pronotum; aedeagus with broad weakly-sclerotized dorsal medial flap.

Argentina, Brazil, Colombia, Costa Rica, Guatemala, Panama, Paraguay, and Venezuela.

Moldenke (1981) subdivided this subgenus into two groups. Argentinian species belong to type II group, which are characterized by having protibiae twice carinate on posterolateral surface; and antennomere IV 1/3 - 3/4 times width of V.

1a. Lacordaire, 1848 (MNS, CTS, CHA, JUY).

1b. Lacordaire, 1848 (MNS, CTS, CHA).

= Lacordaire, 1848

2. Moldenke, 1981 (MNS).

Moldenke, 1981

Lacordaire, 1848, designated by Moldenke (1981: 113).

Scutellum length 1.5x or more than length of posterior lobe of pronotum; frons of male with deep medial depression; dorsal region of aedeagus with very prominent row of setae, no cleft, ventral lobe absent, apex of aedeagus extraordinarily truncate.

Argentina, Bolivia, Brazil, Paraguay, and Peru.

Moldenke, 1981 (CAT). Host plant: : sp.

(Lacordaire, 1848) (MNS, SAL).

= Lacordaire, 1848.

incertae cedis by Moldenke (1981)

1. Monrós, 1953a (SEO, SFE).

2. Lacordaire, 1848 (sensu Monrós) (JUY, SAL, TUC).

= Lacordaire, 1848.

= Lacordaire, 1848.

As mentioned above, further studies are necessary to clarify the relationships of and with the remaining subtribes.

Chevrolat, 1836

Fig. 16

Figure 16.

Lacordaire (2), left: habitus (dorsal view), right: habitus (lateral view).

Chevrolat in

(Olivier, 1791), designated by Monrós (1953b: 46).

This genus has several diagnostic characters that clearly separate it from all other Neotropical : scutellum inclined posterodorsally, distinctly protruding from the plane of elytra; intercoxal prosternal process nearly absent between fore coxae; epipleural margin broadly angled, tip rounded; elytra without pubescence; frons with medial pit, densely and coarsely punctate throughout; pygidium strongly convex; pronotum with lateral margins sulcate to receive antennae in repose; dorsum brilliant metallic.

From Mexico to Argentina.

1a. Lacordaire, 1848 (CTS). Host plant: ? and curupatí (Monrós, 1953a).

1b. Lacordaire, 1848 (CHA, CTS, ERS, FOR, JUY, MNS, SAL, SEO, SFE, TUC). Host plants: : sp.; sp.; sp.sp. (Monrós, 1953a). : sp. (Naranjo) (Monrós, 1953a).

Lacordaire, 1848.

2. Lacordaire, 1848 (TUC, SAL).

= Monrós, 1953a.

Major classification changes in were implemented based on the development of sexual dimorphic characters, especially as they relate to head modifications in males (Agrain and Roig-Juñent 2011). The monophyly of the subtribe is well supported by a set of synapomorphies, including external morphology and genitalia (Agrain and Roig-Juñent 2011).

Lacordaire, 1848

Fig. 17

Figure 17.

(Monrós) (1), left: habitus (dorsal view), right: habitus (lateral view).

Lacordaire, 1848, designated by Monrós (1953a: 114).

This genus can be separated from by the lack of bilobed lacinia and the elytra with confused punctation, in some cases exceedingly coarsely and deeply punctate. Other useful diagnostic characters are: head moderately prominent, covered with dense fine punctation and silky pubescence; male head as long as wide; eyes round and salient; prothorax transverse, pronotal disc as high as long; scutellum often coarsely punctate and always with dense white pubescent; elytra either metallic unicolored and glabrous or black; ventrites usually covered with exceedingly dense white pubescence; female anal pit normally small and only moderately depressed.

USA to Argentina.

As for other groups within , it is in need of modern taxonomic revision. Several species groups have been proposed, but their monophyly has not yet been tested. Monrós (1953a), split this genus into two informal species groups, mainly based on sexual dimorphism evident in male heads. Moldenke (1970) proposed six informal species groups. Subsequently, Moldenke (1981) preserved only three of his earlier groups and transferred some species to two new genera ( and ). More recently, Agrain and Roig-Juñent (2011), recovered a monophyletic clade (sister to ), containing the type species of , , and . This clade is supported by two synapomorphies: male head as long as wide, and pronotal disc as high as long. Consequently, the latter two genera were synonymized with . Some North American species are found in the nests of ant genera Mayr and Linnaeus. Moldenke (1981) divided this genus into three species groups and he indicated three species to be present in Argentina: Monrós, and Lacordaire within group II; and Harold within group III.

1. (Monrós, 1953a) (BAS, CHA, COR, ERS, MNS, SEO, SFE). Host plant: : sp. (branches). : on flowers (Monrós, 1947).

= Guérin, 1945; nec Lacordaire, 1848.

Monrós, 1953a.

Moldenke, 1981.

2. Burmeister, 1877 (COR, ERS, SEO).

Euroscopa (: Moldenke, 1981 (misspelled for ).

3. Monrós, 1953a (MNS).

Euroscopa (: Moldenke, 1981 (misspelled for ).

4. Monrós, 1953a (CHA, FOR).

Euroscopa (: Moldenke, 1981 (misspelled for ).

5. Guérin, 1949 (COR, MZA, SEO).

= Guérin, 1944, not Burmeister, 1877.

Euroscopa (: Moldenke, 1981 (misspelled for ).

6. Monrós, 1953a (SAL, SEO).

7. (Monrós, 1953a) (CHA, CTS, FOR, MNS).

: Moldenke, 1981.

Monrós, 1953a.

8. Monrós, 1953a (CAT, CHA, COR, MZA, SAL) Host plant: : sp. (Roig-Juñent, 2004).

Euroscopa (: Moldenke, 1981 (misspelled for ).

9. Guérin, 1945 (COR, SEO).

Euroscopa (: Moldenke, 1981 (misspelled for ).

10. Lacordaire, 1848 (CTS).

Lacordaire, 1848 (incertae sedis in Moldenke, 1981).

10. Lacordaire, 1848 (CTS, MNS).

: Monrós, 1953a.

11. Harold, 1875 (CHA, COR, ERS, FOR, MZA, SAL TUC, SEO). Host plant: : (Mol.) Mol. (Monrós, 1953a).

Lacordaire, 1848

Lacordaire, 1848: 493–494;

Moldenke (1981) divided this genus into three subgenera, one of these, E. (, was synonymized by Agrain and Roig-Juñent (2011) with . From the remaining two subgenera only one has representative species in Argentina, .

Moldenke, 1981

Fig. 18

Figure 18.

Lacordaire (1), left: habitus (dorsal view), right: habitus (lateral view).

Lacordaire, 1848, designated by Moldenke 1981: 94.

Size greater than 7mm; robust, widest at humeral angle; scutellum foveate; aedeagus occupying entire abdominal length, distinctly straight; body not metallic, eyes weakly emarginate; antennomere IV smaller than V; elytral punctation seriate or sub-seriate, without pubescence; pygidium with transverse subapical carina.

Argentina, Brazil, Colombia, Costa Rica, French Guiana, Mexico, Peru, Suriname, and Venezuela.

1. Lacordaire, 1848 (MNS).

Lacordaire, 1848.

: Moldenke, 1981.

2. Lacordaire 1848 (COR, MNS). Host plant: :

sp. (Monrós 1947).

(Moldenke 1981) Lacordaire, 1848.

: Moldenke, 1981.

Chevrolat, 1836

Fig. 19

Figure 19.

(Forsberg) (1), left: habitus (dorsal view), right: habitus (lateral view).

(

(Germar, 1824) [= (Forsberg 1821)], subsequent designation by Monrós 1953b: 46.

Among the species of , several morphological differences exist and the head and thorax are highly variable, therefore, the most useful PageBreakmorphological characters are: presence of a carina in the inter-ocular area, development of anterior teeth on the mandibles, clypeus sculpture, and the degree of retraction of the head inside the prothorax. The thorax may have strong constrictions, which are often present in those species showing great development of the head and mouthparts. The elytra are also variable; the most distinctive characters are the coloration pattern and the ordering of the elytral punctation. Although also variable, the abdomen and legs are not especially useful for the recognition of species groups. The pygidium may possess distinct sculpture patterns, which are useful to diagnose among species.

distribution includes North, Central and South America, especially diverse in xeric temperate or subtropical zones.

This genus was revised by Agrain (2013). now includes 43 species (Agrain 2014), 13 of them present in Argentina. According to Agrain and Roig-Juñent (2011) this genus is supported by two synapomorphies: eyes strongly emarginated and dorsal plate of aedeagus with straight margin. The morphology in the genus is highly variable, especially for sexual dimorphic features such as: a greater development of mandibles, the presence of constrictions on the pronotal disc, development of double infraocular projection and lengthening of forelegs (Agrain 2013).

1. (Forsberg, 1821) (COR, CTS, FOR, MNS, SEO).

= Forsberg, 1821.

= Germar, 1824.

= Germar 1824.

: Lacordaire, 1848.

= Achard, 1926.

= Achard, 1926.

2. Achard, 1926 (CAT, CHA, COR, CTS, FOR, JUY, LRA, MZA, SAL, SEO, SLS, TUC). Host plants: : (Roig-Juñent 2004), (Roig-Juñent 2004), sp. (Viana and Williner 1974).

: Agrain et al., 2007.

= Monrós, 1945 (nec Lacordaire, 1848).

: Monrós 1956a.

3. Lacordaire, 1848 (COR, SAL, SEO).

= Dejean, 1836 (nomen nudum).

Lacordaire, 1848.

= Achard, 1926.

= Achard 1926.

= Guérin, 1949.

4. Lacordaire, 1848 (CAT, CHA, COR, CTS, ERS, FOR, JUY, LPA, LRA, MNS, MZA, SAL, SEO, SFE, SJN, SLS, TUC). Host plants: : sp.; sp. (Monrós 1953a); (Aravena PageBreak1974), (Roig-Juñent 2004). Ant hosts: Colonies of sp. (Monrós 1953a, as Dr. Oblobin pers. comm.)

Lacordaire, 1848.

= Achard, 1926.

= : Monrós, 1953a (misspelling pro clavapex).

= Achard, 1926.

= Achard, 1926.

= Lacordaire, 1848.

= Lacordaire, 1848.

5. (Forsberg, 1821) (COR, CTS, FOR, JUY, MNS) .

= Forsberg, 1821.

= Germar, 1824.

= : Dejean, 1836.

= Dejean, 1836 (nomen nudum).

Lacordaire, 1848.

= Achard, 1926.

= Achard, 1926.

= Achard, 1926.

= Achard, 1926.

6. Lacordaire, 1848 (COR, MNS).

Lacordaire, 1848.

7. Lacordaire, 1848 (CAT, CHA, CTS, FOR, LRA, MNS, MZA, SAL, SEO, SFE, SJN, SLS, TUC). Host plants: (Monrós 1953a) (Brea); sp.; (Chañar) (Viana and Williner 1974); (Mol.) Mol. (Viana and Williner 1974); (Agrain and Marvaldi 2009); sp. and (Cordo and DeLoach 1995): . : (Common name in Argentina: Retamo, in Peru: Calato).

= Dejean, 1836 (nomen nudum).

Lacordaire, 1848.

= Harold, 1875.

= Achard, 1926.

= Achard, 1926.

= Achard, 1926.

= Achard, 1926.

= Achard, 1926.

= Achard, 1926.

= Achard, 1926.

= Achard, 1926.

8. Lacordaire, 1848 (CTS, ERS) Host plants: : (Guacatonga or wild coffee) (Agrain 2013).

Lacordaire, 1848.

= Lacordaire, 1848.

= Monrós, 1953a.

9. Lacordaire, 1848 (CTS, MNS).

= Dejean, 1836 (nomen nudum).

Lacordaire, 1848.

= Lacordaire, 1848.

10. Monrós, 1953a (MNS).

= Monrós, 1953a.

11. (Germar, 1824) (CTS, MNS).

= Germar, 1824.

: Lacordaire, 1848.

= Lacordaire, 1848.

= Monrós, 1945 (nec Lacordaire, 1848).

= Monrós, 1953a.

12. Lacordaire, 1848 (MNS, SAL).

= Lacordaire, 1848.

= Monrós, 1953a.

13. Monrós, 1947 (MNS).

Monrós, 1947.

Lacordaire, 1848

Fig. 20

Figure 20.

Baly (1), left: habitus (dorsal view), right: habitus (lateral view).

(Germar, 1824), designated by Monrós 1953a: 130.

This genus is close to , from which it can be separated by its unicolored metallic green/blue coloration (except in and , not present in Argentina) and by its distinct body shape. Eyes large, non-emarginate and distinctly protruding; elytra glabrous without distinct punctation pattern, epipleurae reduced; pygidium flat; antennomere IV much smaller than V, not clavate; ventrites densely pubescent.

From Costa Rica to Colombia, and in Brazil, Paraguay, and Argentina.

1. Baly, 1877a (CTS, MNS, TUC).

Baly, 1877a.

Tribe Gyllenhal, 1813

The most common characters to differentiate this tribe are the procoxae globose, distinctly separated by intercoxal prosternal process. In males of many species of () and () the dorsal lobes of the eyes are strongly converging towards the median line and may come into contact with each other. Phylogenetic significance (if any) of this trait is unclear. The antennae are long and filiform in most genera, often reaching the humeral callus or further, although there are exceptions (eg. clavate in ).

Subtribe Gyllenhal, 1813

The most distinctive character is the crenulate, not margined, base of pronotum. Some characters present variation, such as the tarsal claws which may be simple or appendiculate, or antennae, which may be short and clavate to subserrate.

Geoffroy, 1762

Fig. 21

Figure 21.

Burmeister (2), left: habitus (dorsal view), right: habitus (lateral view).

Geoffroy, 1762: 231 (conserved name);

Müller, 1764 (subsequent use).

Chevrolat, 1836.

Chevrolat, 1836 (nomen nudum).

Saunders, 1842.

Saunders, 1842.

Saunders, 1843.

Saunders, 1845, not

Saunders, 1845.

: Saunders, 1845 (error).

Haldeman, 1849.

Stål, 1857.

Baly, 1877c.

Type species.

, designated by Latreille 1810: 432.

Diagnosis.

Anteriorly flat head, deeply inserted into the prothorax; eyes reniform; leading edge of prothorax laterally straight; denticles present on posterior margin of pronotum; thorax closely fitted to base of elytra (thus sometimes concealing denticles); anterior margin of intercoxal prosternal process uniformly concave or with medial flange; intercoxal width equal to or greater than width of coxal cavity; ventrite I of male without spines. Rectal apparatus bearing one ventral and two dorsal sclerites.

Distribution.

Worldwide, with over 1700 species (Chamorro 2014b), with nine species cited for Argentina.

Remarks.

Although a complex subgeneric classification does exist for Palearctic species (Schöller 2002), new world species including Argentinian, have not yet been assigned to subgenera.

Argentinian species checklist.

1. Jacoby, 1907 (TUC).

2. Jacoby, 1907 (BAS).

3. Burmeister, 1877 (BAS).

4. Suffrian, 1863 (BAS).

5. Suffrian, 1863 (BAS).

6. Suffrian, 1857 (BAS, CTS).

7. Jacoby, 1907 (TUC).

8. Jacoby, 1907 (TUC).

Subtribe Leng, 1920

The members of this subtribe have the intercoxal prosternal process noticeably wider than long; tarsal claws appendiculate; antennae are usually short (rarely longer than base of pronotum) and antennomeres expanded laterally.

Gistel, 1848

Fig. 22

Figure 22.

(Suffrian) (2), left: habitus (dorsal view), right: habitus (lateral view).

Gistel, 1848: 123;

Fabricius [= (Fabricius)], designated by Balsbaugh (1966: 660).

may be most commonly confused with and less so with . Both genera in have shorter antennae (rarely surpassing half of entire body length) with antennomeres anteriorly expanded (less so in ). Also, the intercoxal prosternal process is wide and bilobed with small lateral projections and the anterior margin of intercoxal prosternal process uniformly concave (Chamorro-Lacayo and Konstantinov 2004). can be distinguished from by the more uniform rounded shape of the pronotum, which lacks basolateral depressions. The prothoracic anterior opening in has a circumference, best viewed anteriorly, with the dorsal and ventral margins on the similar vertical plane (in lateral view). In general, is larger, more robust, and colorful than .

This genus is restricted to the New World, from México to Argentina with over 100 species. 11 of which are present in Argentina.

Balsbaugh (1966) revised the North American species of this genus. The limits of the subtribe, genera, and species need revision. Information is lacking for Central and South American species that are known only from their original descriptions in the 19th century. The presence of denticles on the posterior margin of pronotum is shared with , therefore may not be a natural group, and may be a synonym of . This hypothesis remains to be tested.

1. (Burmeister, 1877) (Patagonia, RNO).

2. (Boheman, 1858) (BAS, CTS).

3. (Suffrian, 1863) (TUC).

4. (Suffrian, 1863) (BAS).

5. (Burmeister, 1877) (SCZ).

6. (Burmeister, 1877) (Patagonia, SCZ).

7. (Suffrian, 1863) (Argentina).

8. (Boheman, 1858) (Argentina).

9. (Boheman, 1858) (BAS).

10. (Jacoby, 1908) (TUC).

11. (Burmeister, 1877) (BAS).

Baly, 1877

Fig. 23

Figure 23.

(Suffrian) (2), left: habitus (dorsal view), right: habitus (lateral view).

Suffrian, by original designation.

Coxa widely separated, epipleural lobes strongly produced. Chamorro-Lacayo and Konstantinov (2004), provided several prothoracic characters: pronotal punctures absent; intercoxal prosternal process truncate; anterior margin of intercoxal prosternal process with a medial flange, or two submedial flanges.

From Costa Rica to Argentina, mainly in tropical regions.

Weise (1921) disagreed with the separation of this genus from . White (1968), interpreted Weise’s comment as the synonymyzation of with . Since Weise (1921) only provided morphological differences of Suffrian, which is not the type species of the genus, we still consider as a valid genus. Chamorro is currently revising the genus.

Baly, 1877c (MNS).

(Suffrian, 1863) (TUC). This species newly cited for Argentina (Tucuman, Famaillá: Quebrada de Lules, 30-I-1942 // F. Monrós Collection 1959).

Subtribe Chapuis, 1874

The following characters (when combined) can help with the identification of its members (Chamorro 2013): Presence of tibial spurs (absent in , , and s. str.); lack of denticles on the posterior margin of the pronotum (i.e. not crenulate); base of pronotum margined and bilobed sinuate (except in and less or differently margined in ); coarsely punctate dorsally and ventrally including hypomeron (except , and ); confused elytral punctures (except ; less orderly in , ); intercoxal prosternal process lobed (bilobed in other groups) and posterior margin produced caudad (less so in ; eyes visible from above (not visible from above in and ; bulging, particularly in , and , in all other genera the dorsal section of the eye is generally larger than the ventral part as separated by the well developed canthus (canthus weak in , and . This subtribe is currently being revised by Davide Sassi.

Haldeman, 1849

Fig. 24

Figure 24.

(Suffrian) (4), left: habitus (dorsal view), right: habitus (lateral view).

Haldeman, 1849: 245.

Suffrian, 1852.

Haldeman 1849: 245 (= Fabricius, 1801), by monotypy.

Recently, Chamorro (2013) provided the following characters to identify the species of this genus: posterior margin of intercoxal prosternal process rounded; lateral margin of elytra deeply excised exposing abdomen caudally, elytra length approximately 2× or less length of pronotum. However, the limits of and are confused and require revision.

Nearctic and Neotropical.

(Suffrian, 1866) (BAS, COR).

(Suffrian, 1866) (BAS).

(Suffrian, 1866) (BAS).

(Suffrian, 1866) (BAS).

Burmeister, 1877 (ERS).

Burmeister, 1877 (BAS).

Suffrian, 1866

Fig. 25

Figure 25.

Jacoby (4), left: habitus (dorsal view), right: habitus (lateral view).

Not yet designated.

includes species that lack a deeply excised lateral edge of the elytra, additionally, the following characters may be useful to segregate species PageBreakinto this genus: posterior margin of intercoxal prosternal process gradually narrowing, pointed; abdomen not exposed; elytral length greater than 2× length of pronotum (Chamorro 2013).

Neotropical.

This genus has not been revised since its original description by Suffrian (1866) and its relationship with related genera is presently unclear (Schöller 2003, Sassi 2015). Furthermore, Jacoby (1907) indicated and to be very problematic to define, and a lot of species can not fit well in either genera. However, a study with a new diagnosis of the genus , based on a new set of effective morphological characters is in progress (Sassi, in prep.).

1. Suffrian, 1866 (CTS).

2. Suffrian, 1866 (BAS, COR).

3. Jacoby, 1907 (COR).

4. (Jacoby, 1907), comb. n. (BAS).

5. Jacoby, 1907 (SFE).

6. Suffrian, 1866 (CTS).

7. Suffrian, 1866 (BAS, CTS).

8. Suffrian, 1866 (Argentina).

9. Suffrian, 1866 (COR, SLS). Host plant: : carqueja ( sp.) (Viana and Williner 1973).

10. Jacoby, 1907 (SFE).

11. Suffrian, 1866 (BAS, COR, MZA, RNO).

12. Suffrian, 1866 (CTS).

Stål, 1857

Fig. 26

Figure 26.

(Suffrian) (4), left: habitus (dorsal view), right: habitus (lateral view).

:

Stål (= Blanchard), designated by Jakobson 1924: 258.

This genus can be easily distinguished from all others in the area by the presence of pubescence on its body and by the presence of a basal thoracic lobe with raised, thickened apex. Furthermore, it has nearly entire eyes and the rectal apparatus bears two ventral and three dorsal sclerites, with the shape of the dorsal central plate band-like (very narrow) (Reid 1990; Schöller 2008).

This genus has eight species described form Southern Chile and Argentina, and some new species awaiting description. Species are found between 30°S and 42°S and are associated with sclerophyllous shrubs (Jerez and Briones 2010).

This genus was considered a synonym of by several authors, however several studies support its validity and it is hypothesized to be included in (Baly 1877b; Jakobson 1924; Monrós 1949a, Schöller 2008; Jerez and Briones 2010) or in its own subtribe (Reid 1990; Chamorro and Konstantinov, unpublished data).

1. Monrós, 1949a (CHU, RNO). Host plant: : (Monrós 1949a).

2. (Blanchard, 1851) (NQN, RNO). Host plant: : “litrenillo”, ? (Bosq 1943), sp. (Monrós 1949a); : sp., : sp., and , (as main food source) (Monrós 1949a), : sp. (Monrós 1949a), sp. (Bosq 1943), : sp. (Monrós 1949a).

= Stål, 1857.

3. Monrós, 1949a (RNO). Host plant: : sp. (Monrós 1949a).

4. Monrós, 1949a (NQN).

5. (Suffrian, 1863) (NQN, RNO).

6. (Suffrian, 1866) (NQN, RNO, CHU). Host plant: : (Monrós 1949a).

Chevrolat, 1836

Fig. 27

Figure 27.

Suffrian (2), left: habitus (dorsal view), right: habitus (lateral view).

: Chevrolat, 1836: 420.

Redtenbacher, 1845.

Rey, 1883.

:

Laicharting, 1781. By subsequent designation of Jacoby 1908: 265.

Posterior margin of intercoxal prosternal process relatively entire, rarely produced beyond posterior margin of prothorax; gestalt cylindrical (height of each elytron approximately 2.5 width), pronotum narrower than elytral bases combined, overall flattened not vaulted; punctation on head, prothorax and elytra evident, large; elytral punctation commonly confused (but punctation in rows not uncommon); fore-femora may or may not be enlarged; mesotibiae usually with terminal spur in both sexes.

Neartic, Neotropical, Palearctic, and Oriental regions.

A subgeneric classification exists for Palearctic species, Neotropical species have not yet been assigned to subgenera.

1. Suffrian, 1866 (BAS)

2. Blanchard, 1851 (ARGENTINA). Host plant: : and (Jolivet, 1978).

3. Suffrian 1866 (BAS, LPA). Host plant: : (Aravena 1974).

4. Boheman, 1858 (BAS)

5. Suffrian, 1866 (CTS, ERS).

Tribe Jakobson, 1924

= Gressitt, 1946

This tribe is the least diverse within , and it is much more diverse in the Neotropics (Chamorro 2014b). Most adults resemble caterpillar droppings. Chamorro-Lacayo and Konstantinov (2009), undertook a comprehensive synoptic study of the world genera of .

Monrós, 1951c

Fig. 28

Figure 28.

(Lacordaire) (2), left: habitus (dorsal view), right: habitus (lateral view).

Monrós, 1951c: 657;

Lacordaire, 1848, by original designation.

The most salient feature of this genus is the presence medially on the pronotum of six elevated distinct, small, sharp, longitudinal carinae, which converge medially near the posterior margin, reminiscent of a fan. These are small beetles (less than 3 mm length), cylindrical; with antennae serrated beyond antennomere V. antennomeres III-V slightly widened, but not dilated distally; pronotal base opposite mesoscutellum (posterior pronotal lobe) with or without notch; intercoxal prosternal process gradually narrowing posteriorly, broadening before apex; metascutellum concealed by elytra; elytral suture completely serrate, although serration may be weak near scutellum, elytral tubercles well developed. Tibiae slightly curved, cylindrical. can easily be distinguished from Rafinesque by the presence of the six longitudinal carinae on its pronotum.

Pantropical, except Australia (Monrós 1951c). Six of the 21 Neotropical species are present in Argentina.

1. (Lacordaire, 1848) (CTS, JUY, MNS).

2. Monrós, 1951c (MNS).

3. Monrós, 1951c (MNS).

4. Monrós, 1951c (MNS).

5. Monrós, 1951c (CTS, MNS, TUC).

6. Monrós, 1951c (COR).

Rafinesque, 1815

Fig. 29

Figure 29.

(Lacordaire) (3), left: habitus (dorsal view), right: habitus (lateral view).

Rafinesque, 1815: 116;

Knoch, 1801: 122.

Ihering, 1907.

Ihering, 1904.

Jakobson, 1924.

Knoch, 1801, by subsequent designation of Navajas 1946: 244 [preoccupied by Röding, 1798, ].

This genus can be separated from Lacordaire by the following characters: males without spines or spinulae on ventrite I; antennomere V nearly as large as VI; sutural serration of elytra usually incomplete (suture entire immediately following mesoscutellum); intercoxal prosternal process posteriorly pointed (narrowed), posteriorly much narrower than anterior margin (Chamorro-Lacayo and Konstantinov 2009). Intermediate size (3-8 mm length). Body usually not metallic in color; elytra without velvety spots. Antenna serrate beyond antennomeres III or IV, antennomere II slightly widened, globose, antennomere V nearly as large as 6th. Pronotum medially elevated, with various bumps and short carinae; posterior pronotal lobe with well-differentiated notch; metascutellum not exposed.

Cosmopolitan, with over 400 species described worldwide (Monrós 1951c; Reid 1991).

1. Monrós, 1951c (LRA, SAL).

2. Monrós, 1951c (JUY, SAL, TUC).

3. (Lacordaire, 1845) (CHT, NQN, RNO).

= Blanchard, 1851.

= Philippi & Philippi, 1864.

= Philippi & Philippi, 1864.

4. Monrós, 1951c (MNS).

5. Monrós, 1951c (JUY).

6. Monrós, 1951c (CHA).

7. (Jacoby, 1901) (MNS). Host plant: : (Bokermann 1963).

8. (Klug, 1824) (SAL) Host plant: : (Bokermann 1963).

9. (Lacordaire, 1848) (BAS, CHA, COR, ERS, FOR, JUY, MNS, SAL, TUC)

= Lacordaire, 1848. Host plant: : (Bokermann 1963).

10. Monrós, 1951c (CTS).

11. (Kollar, 1824) (CTS, MNS). Host plants: , , , , , (Monrós 1951c).

12a. (Klug, 1824) (BAS, CHA, COR, FOR, JUY, LRA, MNS, SAL, SEO, SFE, SLS, TUC).

= Jacoby, 1901. Host plants: : sp., ; : “Tasi” (Monrós 1951c).

12b. Monrós, 1951c (COR, LRA, MZA, SEO).

13. (Fabricius, 1801) (MNS).

14. Monrós, 1951c (CTS).

15. Monrós, 1951c (MNS).

16. (Kollar, 1824) (MNS).

17. Monrós, 1951c (JUY, SAL).

18. (Kollar, 1824) (MNS).

= Klug, 1824. Host plant: : (Bokermann, 1963).

19. Monrós, 1951c (MNS).

20. Monrós, 1951c (COR, ERS, SAL, TUC). Host plant: : sp., : sp. (Monrós, 1951c), (Bokermann 1963).

21. Monrós, 1950 (BAS, COR, CTS). Host plant: : sp. (Bokermann 1963).

22. (Kollar, 1824) (FOR).

= Klug, 1824.

23. Monrós, 1951c (FOR). Host plant: : sp. (Monrós 1951c).

24. Monrós, 1951c (MNS).

25. Monrós, 1951c (MNS).

26. Monrós, 1951c (JUY).

27a. (Germar, 1824) (JUY).

= Kollar, 1824. Host plant: : sp. (Monrós, 1951c).

27b. (Lacordaire, 1848) (COR, MNS).

28. (Lacordaire, 1848) (CTS).

= Lacordaire 1848. Host plant: : , and (Bokermann 1963).

29. (Lacordaire, 1848) (MNS, SAL).

30. Monrós, 1951c (CHA, COR, CTS, FOR, JUY, MNS, SAL, TUC). Host plant: : (Monrós, 1951c).

31. Monrós, 1951c (CHA, CTS, FOR, JUY, MNS, SAL, SFE, TUC).

32. (Kollar, 1824) (MNS).

= Klug, 1824. Host plant: : (Bokermann 1963).

33. Monrós, 1951c (JUY, SAL, TUC). Host plant: : sp. (Monrós 1951c).

34. Monrós, 1951c (MNS).

Lacordaire, 1848

Fig. 30

Figure 30.

Monrós (2), left: habitus (dorsal view), right: habitus (lateral view).

Lacordaire, 1848: 844;

Kollar, 1824, by subsequent designation.

Small species (2–3.5 mm length), cylindrical with widest near shoulders; antenna serrate beyond antennomere V, antennomeres III-IV slightly widened, but not dilated distally; pronotum with various bumps and short ridges, posterior pronotal lobe concave, usually without well differentiated notch.

Present in Nearctic, Neotropical, and Oriental regions (Monrós 1951c; Karren 1966). Includes 26 species, 10 represented in the Neotropics.

Gressitt and Kimoto (1961) synonymized this genus with , yet, this decision has been ignored and is considered to be a valid genus (Karren 1966, 1972; Seeno and Wilcox 1982; Riley et al., 2003). The relationship among genera remains to be studied.

1. Monrós, 1951c (COR, MNS).

2. (Kollar, 1824) (MNS).

= Kollar, 1824.

= Klug, 1824 (nec Kollar, 1824). Host plant: (Monrós 1951c).

3. Monrós, 1951c (CHA, FOR, SAL, TUC). Host plant: : sp. (Jolivet 1978).

4. Monrós, 1951c (COR, JUY, MNS, SAL, SLS). Host plant: : and (Bokermann 1963).

Voet, 1806

Fig. 31

Figure 31.

(Kirby) (2), left: habitus (dorsal view), right: habitus (lateral view).

Voet, 1806 = Fabricius, 1798, by monotypy.

This genus includes some of the larger and more charismatic species in the group (6.5–7.2 mm length). Body subquadrate and metallic, antenna serrate beyond antennomere III; the head with a longitudinally impressed vertex; elytral tubercles pronounced; posterior pronotal lobe with an acute notch; sutural serration of elytra well-developed beyond the middle of suture towards apex; ventrite I with lateral tubercles; fore- and midtibial apices with spine; tarsal claws simple. According to Chamorro-Lacayo and Konstantinov (2009), can be distinguished from all other genera of the tribe by the longitudinally impressed vertex of the head, simple tarsal claws, large body size, and usually bright metallic coloration.

From Mexico to Argentina, with seven species.

This is a small genus with only seven described species (Monrós 1951c). Chamorro-Lacayo and Konstantinov (2009), mistakenly cited (Lacordaire) for Argentina, this is present in Goiaz state of Brazil.

1. (Kirby, 1818) (CTS, JUY, MNS, SAL, TUC). Host plants: : sp., sp. (Monrós 1951c); : , , (Bokermann 1963).

Monrós, 1948a

Fig. 32

Figure 32.

(Klug) (2), left: habitus (dorsal view), right: habitus (lateral view).

Monrós, 1948a: 192;

Klug 1824, by original designation.

can be separated from all other genera by the nearly rectangular prosternal process; since the process is more or less triangular in all other genera of warty leaf beetles. Intermediate size (length 3.60-5.20 mm); body shape subglobular; antenna serrate beyond antennomere III, antennomere III slightly dilated distally; pronotum without median elevation, relatively smooth; sutural serration of elytra absent or weakly developed; elytra without well developed tubercles; appendiculate tarsal claws.

The genus include 13 Neotropical species (Chamorro-Lacayo and Konstantinov 2009).

1. (Klug, 1824) (MNS). Host plant: : y sp. (Araça) (Bokermann 1963).

Lacordaire, 1848

Fig. 33

Figure 33.

(Jacoby) (2), left: habitus (dorsal view), right: habitus (lateral view).

Lacordaire 1848, by monotypy.

can be distinguished from all other genera in the tribe by: head not completely retracted into prothorax; mandibles enlarged in males (sexual dimorphism); intercoxal prosternal process strongly and abruptly constricted beyond anterior margin; and prosternal process more than ¾ as long as intercoxal prosternal process. These beetles are small sized (length 3.45-4.72 mm), cylindrical; body usually yellowish; canthus of eye as yellow as rest of frons; pronotum and elytra glabrous; head not completely retracted into prothorax; mandibles enlarged in males; antenna serrate beyond antennomere III, antennomere II slightly widened, globose, antennomere V as large as VI; posterior pronotal lobe with well differentiated notch; intercoxal prosternal process strongly and abruptly constricted beyond anterior margin; sutural serration of elytra complete; elytral tubercles poorly developed; tarsal claws bifid or appendiculate.

This genus contains only five species, distributed in North, Central, and South America (Chamorro-Lacayo and Konstantinov 2009; Karren 1972).

(Jacoby, 1904) (MNS).

LACORDAIRE, 1848

Adults: Body compact, strongly convex; head inserted into prothorax (not visible from above). Pronotum convex tightly appressed to elytral base; antennal groove present on each side of prosternal process. Elytra covering pygidium. Larva differs from as pointed out in previous section.

Tribe Lacordaire, 1848

This tribe is composed of 10 genera (Seeno and Wilcox 1982) and 250 species (Chamorro 2014a). Four genera occur in the Neotropical region (Chamorro 2014a): Lacordaire, Monrós, Kirby, and Monrós. It is the only Lamprosomatine tribe represented in Argentina where the fauna comprises 1 genus, , and 8 species. is characterized by the presence of a file on distal margin of last ventrite; last ventrite not excised in shape of arc; pygidium completely covered by elytra; scutellum acutely triangular (small to very small); elytral punctuation arranged in regular rows or with a tendency to form such rows.

Kirby, 1818

Fig. 34

Figure 34.

Germar (3)., left: habitus (dorsal view), right: habitus (lateral view).

Kirby, 1818: 445. By monotypy.

body length about 4.5 mm; tarsal claws appendiculate with broad tooth; antenna short, antennomere VIII nearly as wide as VII or IX. Metallic coloration (some species multicolored), head not visible from above, clypeus excavate. According to Monrós (1956b) it can be differentiated from other Neotropical genera by having appendiculate claws at 180º angle, while and have simple claws at a more obtuse angle.

Nearctic, Neotropical, in Argentina limited to north, and northeastern provinces.

Adults feed on plants of the families , , , . Therefore, some species have been considered as potential biological control agents for these plants (Caxambú and Almeida 2003).

1. Monrós, 1948b (JUY, SAL, TUC). Host plant: : spp. (Bark-gnawing).

2. Germar 1824 (MNS). This species newly cited for Argentina.

3. Monrós, 1948b (CHA).

= Monrós, 1948b. Parasitoids: and () (Monrós 1948b). Host plant: : sp. (Monrós 1948b).

4. Monrós, 1947 (CTS).

5. Monrós, 1948b (SAL). Host plant: : (Bark-gnawing).

6. Monrós, 1948b (CTS).

7. Guérin-Ménéville, 1844 (Northeast Argentina).

8. Monrós, 1947 (MNS).

Discussion and conclusions

This is the first comprehensive synthesis of Argentinian Camptosomata. This study may prove useful also for countries bordering Argentina. Similar contributions indicated the diversity of Camptosomata in other Neotropical countries as follows: Maes (1998) recorded 19 genera and 46 species for Nicaragua, Chaboo and Schöller (2016) accounted for 14 genera and 43 species for Peru; and in Brazil, 723 species, 26 subspecies in 37 genera of (Sekerka et al. 2015) and 62 species in 5 genera of (Sekerka 2017) were recorded.

Species richness and distribution patterns

Historically, Argentina has been divided in two main regions: Andean and Neotropical (Morrone, 2014). As depicted in the distribution pattern of Camptosomata tribes and subtribes by province (Fig. 35A–D), tribes are mostly distributed in the Neotropical region, while few species reach the Andean region or are found below 40º S latitude. Based on the map of species richness by province (Fig. 36) higher richness (up to 80 species) roughly coincides with the line dividing the Neotropical and Andean regions (Figs 1, 37). Poor data notwithstanding, this pattern fits at the latidudinal diversity graPageBreakPageBreakdients hypothesis with greater species richness at tropical latitudes (Hillebrand 2004). Most of the central and northern provinces (e.g. CHA, COR, CTS, FOR, MNS, SAL, SEO, TUC) are presented on the left side of Figure 37, while most of southern provinces (CHT, CHU, NQN, SCZ, STZ, and TFO) appear on the right side, with few exception, for example the lack of information for SJN, ERS, or SLS.

Figure 35.

A Administrative divisions representing presence of subtribes of as indicated in reference B Administrative divisions indicating presence of subtribes of as indicated in reference.

Figure 36.

Map indicating species number (0–80) (richness) by province.

Figure 37.

Bars diagram showing species by provinces, it can be observed that species richness diminished through southern provinces.

Within (Fig. 35A), is present in the northeastern provinces (FOR, MNS, SEO), while is distributed throughout most of northeastern Argentina. is present from northern Argentina to the central PageBreakregion (as far as MZA, LPA, and BAS). covers this same region, yet it reaches the southern provinces (NQN, and RNO). has not been reported for the southern provinces beyond Rio Negro.

, on the other hand, is putatively mostly absent from the Northwestern provinces of Argentina. The presence of this tribe in Tucumán might indicate a more widespread distribution. Sampling bias and poor inventory may explain the absence of in central and western regions of Argentina. The subtribes of (Fig. 35B) show a more widespread distribution for and , while are mostly recorded from Northeastern Argentina. Finally, seems restricted to the Northern provinces, its absence in Formosa seems artificial, so presence of this subfamily surely will expand with more collecting in this region. According to current information, most species are distributed in the Neotropical provinces, especially: Araucaria forest, Chacoan, Monte, Pampean, Parana Forest, and Prepuna.

Current taxonomic knowledge, basic statistics and future research

A total of 190 species (182 + 8 ) of Camptosomata are currently known from Argentina. The most diverse group of Camptosomata in Argentina is (Fig. 38). However, is also, by far, the most studied group in Argentina due to the efforts of Monrós in the 1950’s. The patchy distribution at administrative division levels clearly indicates the need for specimen identification and incorporation of museum specimens into databases, as well as collection of new specimens. The latter will permit the application of ecological modelling and biogeographic studies of the group that will provide a more complete picture of the biogeographic history and ecological tolerance ranges, as well as help guide conservation policies for the group. The current estimate of endemic species in Argentina is uncertain, and its calculation based on extant information would be inaccurate, especially without a complete species checklist of bordering countries (i.e. Bolivia, Brazil, Chile, Paraguay, and Uruguay). When comparing the timelines in Fig. 39 with the graphic indicating species richness by genus (Fig. 40), it becomes clear that (except for , recently revised by Agrain (2013), several of the most diverse genera have not been revised in over 100 years. Many species are only known from their original descriptions in the mid 19th or mid 20th century (Fig. 39). In many cases, the type specimens were not illustrated. This has resulted in long series of unidentified specimens housed in public and private collections awaiting the study of name bearing types.

Figure 38.

Bars diagram showing the number of genera and species by tribe of Argentinian Camptosomata.

Figure 39.

Timelines showing the years of the last work made on each genera that includes Argentinian species taxonomic treatment (simple checklist included).

Figure 40.

Species number by genus.

Our synthesis here is a necessary step towards further comprehensive study of Argentinian Camptosomata that will facilitate field work, assist in determination of unidentified material housed in South American collections, creation of illustrated of keys to the species level, and with identified specimens in hand achieve databasing of museum specimens. These elementary tasks are prerequisite to modern taxonomic revisions and evolutionary studies.

1	Body dorsally highly convex and ventrally flattened (semicircular in cross-section) (Fig. 2A); abdominal ventrites not connate; antennal grooves present on intercoxal prosternal process; females without a well developed fovea (egg anal pit) on ventrite V (Lamprosomatinae). Distal margin of last ventrite thick (Fig. 2B); last ventrite not excised in shape of arc (Fig. 2C); pygidium completely covered by elytra; scutellum acutely triangular (small to very small); elytral punctation arranged in regular rows or with a tendency to form such rows	Lamprosoma Kirby (Fig. 34). (Lamprosomatini)
–	Body not highly convex, not flattened ventrally (Fig. 2D) (oval in cross-section); abdominal ventrites connate; antennal grooves absent on intercoxal prosternal process (except Fulcidacini and Ischiopachina (Clytrini); females with a distinct, variably shaped fovea (egg anal pit) on ventrite V	2. (Cryptocephalinae)
Cryptocephalinae
2(1)	Intercoxal prosternal process with antennal furrows; body surface usually tuberculate (Fig. 2E); elytral suture usually serrate	4 (Fulcidacini)
2’	Pronotum and intercoxal prosternal process without antennal furrows (Ischiopachina with furrows on hypomeron); body surface not tuberculate; elytral suture entire	3
3(2)	Antennae short (not surpassing the length of pronotum), serrate; procoxae contiguous (Fig. 2F)	9 (Clytrini)
3’	Antennae long (surpassing the length of pronotum, sometimes nearly equal to total body length), filiform, sometimes some segments expanded and flattened; procoxae separated (Fig. 2G)	29 (Cryptocephalini)
Fulcidacini
4(2)	Intercoxal prosternal process nearly rectangular, with posterior margin slightly narrower than anterior margin; pronotal and elytral tubercles reduced, sometimes with velvet patches	Melittochlamys Monrós (Fig. 32)
4’	Intercoxal prosternal process varying in shape, triangular or angulate between mesocoxae, but never rectangular, with posterior margin much narrower than anterior margin; pronotal and elytral tubercles well developed	5
5(4)	Body equal to or greater than 10 mm long; colour brightly metallic; head with vertex longitudinally impressed; tarsal claws simple	Fulcidax Voet (Fig. 31)
5’	Body less than 10 mm long; head with vertex not impressed; tarsal claws usually appendiculate (except Exema (from simple to appendiculate)	6
6(5)	Pronotum with six distinct, small, sharp, longitudinal carinae converging posteromedially, fan-like; color uniform, generally black	Aulacochlamys Monrós (Fig. 28)
6’	Pronotum with or without tubercles, but never with six longitudinal, fan-like carinae	7
7(6)	Head not completely retracted into the prothorax; mandibles in males larger than in females; intercoxal prosternal process strongly and abruptly constricted behind anterior margin; prosternal process more than 3/4 as long as intercoxal prosternal process (Fig. 2H) …Pseudochlamys Lacordaire (Fig. 33)
7’	Head completely retracted into prothorax; mandibles in males as large as in females; intercoxal prosternal process gradually constricted at about 2/3 of its length (Fig. 2I)	8
8(7)	Males without spines or spinulae on ventrite I; antennomere V nearly as long as VI; elytral suture crenulation usually incomplete (i.e., suture entire immediately following scutellum)	Chlamisus Rafinesque (Fig. 29)
8’	Males with spines or spinulae on ventrite I; antennomere V much shorter than VI; elytral suture crenulation always complete	Exema Lacordaire (Fig. 30)
Clytrini
9(3)	Prothorax with lateral antennal grooves on hypomeron Ischiopachina Chapuis (Monotypic subtribe). Elytra without strong parallel longitudinal carinae; color metallic	Ischiopachys Chevrolat (Fig. 16)
9’	Prothorax without lateral antennal grooves on edge of hypomeron	10
10(9)	Tarsal claws simple	12 (Megalostomina Chapuis)
10’	Tarsal claws bifid or appendiculate (Fig. 3A)	11
11(10)	Scutellum flat, in the same horizontal plane as elytra; elytra without longitudinal carinae, frons distinct	15 (Babiina Chapuis)
11’	Scutellum raised above elytral plane; elytra with strong parallel, longitudinal carinae; frons very narrow Arateina Moldenke; (monogeneric subtribe)	Aratea Lacordaire (Fig. 4). (monogeneric subtribe)
12(10)	Eyes entire or only slightly emarginate	13
12’	Eyes strongly emarginate	Megalostomis Chevrolat (Fig. 19)
13(12)	Scutellum foveate; body not metallic; pygidium with transverse subapical carina; aedeagus occupying entire abdominal length	Euryscopa (Coleomonrosa) Moldenke (Fig. 18)
13’.	Scutellum not foveate; body brilliant metallic, or with noticeable metallic reflections; pygidium without subapical transverse carina; aedeagus smaller, not occupying entire abdominal length	14
14(13)	Eyes elongate, not protruding; body robust, subquadrate	Themesia Lacordaire (Fig. 20)
14’	Eyes round and protruding; body elongate	Coscinoptera Lacordaire (Fig. 17)
15(11)	Tarsal claws bifid; dorsum of body uniformly metallic; body lengthened	Helioscopa Gistel (Fig. 9)
15’	Tarsal claws appendiculate; dorsum of body not uniformly metallic; body compact	16
16(15)	Epipleural lobes very pronounced;angle of epipleural lobes rounded; elytra striae strongly impressed; dorsal coloration black, with omnipresent metallic bronze reflections	Saxinis (Saxinis) Lacordaire (Fig. 13)
16’	Epipleural lobes weakly developed, not angulate, no more than two striae wide; elytra striae not strongly impressed; dorsal coloration without bronze reflections	17
17(16)	Pygidium with transverse subapical angle, evenly bent perpendicular to longitudinal axis of body; epipleural lobe not well developed in lateral view	18
17’	Pygidium flat, surface slightly convex, with tip sometimes bent; epipleural lobe distinctly rounded in lateral view	23
18(17)	Eyes feebly emarginate, distinctly projecting, conspicuously protruding at sides of head	Dinophthalma Lacordaire (Fig. 8)
18’	Eyes distinctly emarginate, not markedly projecting	19
19(18)	Anterior margin of pronotum arcuate, entirely covering head from dorsal view; body shape elongate, cylindrical and flat; size large, greater than 10 mm long; frons flat; lateral margins of prothorax not widely explanate	Babia (Coleolacordairei) Moldenke (Fig. 5)
19’	Anterior margin of pronotum transverse or arcuate, but not concealing entire head from dorsal view	20
20(19)	Body shape strongly cylindrical, elongate, not flattened; elytra not fully covering pygidium; elytral punctation barely noticeable	Cylindrodachrys Monrós (Fig. 6)
20’	Body shape not cylindrical or elongate; elytra fully covering pygidium; elytral punctation evident	21
21(20)	Frons strongly tapering, triangular, without transverse sulcus; body shape subquadrate; aedeagus with strong dorsal and ventral tufts of pubescence	Pnesthes Lacordaire (Fig. 11)
21’	Frons not strongly tapering below eyes, subrectangular, with length only slightly greater than width; aedeagus without pronounced ventral and dorsal patches of setae, with only a few dorsal setae present	22. Temnodachrys Monrós (Fig. 14)
22(21)	Frons with deep transverse sulcus (Fig. 3B); body shape subrectangular	Temnodachrys (Temnodachrys) Monrós
22’	Frons without deep transverse sulcus; body guttiform or minute and with subparallel sides	Temnodachrys (Eudachrys) Monrós
23(17)	Body shape subcircular in outline; legs with longitudinal carinae; anterior pronotal margin strongly explanate and completely concealing head from dorsal view	24
23’	Body shape subrectangular, sides subparallel; legs without longitudinal carinae; anterior margin of pronotum never concealing all of head in dorsal view	27
24(23)	Forelegs longer (especially in males) than mid- and hind legs; tarsomere III enlarged, shallowly excavated; head less reflexed, 90º with respect to prosternum (Fig. 3C)	Stereoma Lacordaire (Fig. 12)
24’	All legs with similar development; tarsomere III narrow, deeply excavated; head more reflexed, forming 45º angle with respect to prosternum (Fig. 3D)	25. Urodera Lacordaire (Fig. 15)
25(24)	Posterior margin of pronotum broadly expanded, forming distinct scutellar lobe with angular corners (Fig. 3E)	Urodera (Austrurodera) Moldenke
25’	Posterior margin of pronotum not broadly expanded and not forming a scutellar lobe with angular corners	26
26(25)	Front tibiae with indistinct posterolateral carinae, with surface not deeply excavate and reflective between carinae; frons of male with three shallow depressions	Urodera (Urodera) Lacordaire
26’	Front tibiae with strong posterolateral carinae, with surface deeply excavate and reflective between carinae; frons of male with deep medial depression	Urodera (Stereomoides) Moldenke
27(23)	Pronotum with weak metallic green reflections; antennomere IV much smaller than V; frons wide, with width greater than or subequal to length; frons without medial pit	28. Paraurodera Moldenke. (Fig. 10).
27’	Pronotum without metallic reflections; antennomere IV subequal in size to V; frons narrow, with length more than twice width; frons with deep medial pit	Dachrys Erichson (Fig. 7)
28(27)	Anterior margin of pronotum transverse, not concealing head at all in dorsal view; frons with submedial depressions; sexual dimorphism of frons extreme, the male having extremely wide frons and elongate mandibles	Paraurodera (Torourodera) Moldenke
28’	Anterior margin of pronotum explanate and partially concealing head; frons with medial and two submedial depressions; sexual dimorphism reduced, with frons and mandibles similarly developed in male and female	Paraurodera (Paraurodera) Moldenke
Cryptocephalini
29(3)	Claws simple or, if appendiculate, intercoxal prosternal process longer than wide to subquadrate	30
29’	Claws appendiculate, each with broad, basal tooth; intercoxal prosternal process wider than long	31. Monachulina Leng
30(29)	Pronotum margined at base (except in Mylassa), not crenulate (Fig. 3F)	32 (Pachybrachina Chapuis)
30’	Pronotum not margined at base, usually crenulate (Fig. 3G) (Cryptocephalina Gyllenhal). Eyes with distinct excavation on internal margin; dorsum glabrous; male front tibiae with reduced sexual dimorphism; posterior pronotal margin not produced	Cryptocephalus Geoffroy (Fig. 21)
31(29)	Anterior margin of pronotum simple, arcuate; pronotal punctures distinct throughout; intercoxal prosternal process bilobed, with small lateral projections; anterior margin of prosternum uniformly concave; pronotal anterior opening circular	Lexiphanes Gistel (Fig. 22)
31’	Anterior margin of pronotum produced; pronotal punctures absent; intercoxal prosternal process truncate; anterior margin of prosternum with one or two medial flanges; pronotal anterior opening ventrally widened	Stegnocephala Baly (Fig. 23)
32(30)	Eyes small, bulging, with canthus shallow	33
32’	Eyes large, extending dorsad beyond upper third of head, usually with upper half of eye larger than ventral half; canthus deep, extending approximately 1/4 distance into eye; posterior margin of pronotum (directly opposite scutellum) not produced posteriorly, margined with basal row of punctures, bi-sinuate; scutellum not heart-shaped	34
33(32)	Dorsal surface generally setose; pronotum greatly vaulted, with lateral margins narrow; pronotum medially lobed posteriorly, lobe elevated and truncate; scutellum heart-shaped	Mylassa Stål (Fig. 26)
33’	Dorsal surface glabrous; pronotum regularly convex, with lateral margins prominent, visible from above, with posterior margin regularly biconcave, with mesobasal region regularly rounded and slightly produced posterad; scutellum with posterior margin truncate	Ambrotodes Suffrian (Although this genus has not been yet reported from Argentina, we include it in this key because its species are common along the eastern border of the Andes in Chile.
34(32)	Posterior margin of intercoxal prosternal process convex, produced beyond posterior margin of prothorax; mesotibial spurs present or absent; body robust; punctures not deep or large, particularly on pronotum; dorsal surface shiny	35
34’	Posterior margin of intercoxal prosternal process straight, rarely produced beyond posterior margin of prothorax; gestalt cylindrical (height of each elytron approximately 2.5 width), pronotum narrower than elytral bases combined, overall flattened not vaulted; punctures on head, prothorax and elytra evident, large; elytral punctation commonly confused (but punctation in rows not uncommon); forefemora enlarged or not; each mesotibia usually with terminal spur in both sexes	Pachybrachis Chevrolat (Fig. 27)
35(34)	Posterior margin of intercoxal prosternal process rounded; lateral margin of elytra deeply excised, exposing abdomen caudally; elytra length approximately 2× or less length of pronotum	Griburius Haldeman (Fig. 24)
35’	Posterior margin of intercoxal prosternal process gradually narrowing, pointed; lateral edge of elytra not deeply excised; abdomen not exposed; elytral length greater than 2× length of pronotum	Metallactus Suffrian (Fig. 25)