Cladistic analysis of Hemirhipini with establishment of Propalaus gen. nov. (Coleoptera, Elateridae, agrypninae)

Cladistic analysis of Hemirhipini with establishment of Propalaus gen. nov. (Coleoptera, Elateridae, Agrypninae)

Sônia A. Casari

Museu de Zoologia, Universidade de São Paulo, Caixa Postal 42.494, 04218-970, São Paulo, SP, Brasil. E-mail: [e-mail]

ABSTRACT

This article includes a cladistic analysis of the tribe Hemirhipini. Are included 20 Hemirhipini genera (sensu Casari-Chen 1994), Saltamartinus Casari (1996b) (Hemirhipini), 6 genera excluded from Hemirhipini and kept in Agrypninae (formerly Pyrophorinae) (Casari-Chen 1993) and also, Aphileus Candèze (1857), Pyrophorus Billberg (1820) and Thoramus Sharp (1877). The type-species of the majority of genera and all species of the American genera (except Saltamartinus viduus (Chevrolat 1867)) are included. This analysis demonstrates that 30 genera belong to Hemirhipini: Abiphis Fleutiaux (1926), Alaolacon Candèze (1865), Alaomorphus Hauser (1900), Alaus Eschscholtz (1829), Aliteus Candèze (1857), Anthracalaus Fairmaire (1888), Aphileus Candèze (1857), Austrocalais Neboiss (1967), Calais Castelnau (1836), Catelanus Fleutiaux (1942), Chalcolepidius Eschscholtz (1829), Chalcolepis Candèze (1857), Conobajulus Van Zwaluwenburg (1940), Coryleus Fleutiaux (1942), Cryptalaus Ôhira (1967), Eleuphemus Hyslop (1921), Eumoeus Candèze (1874), Fusimorphus Fleutiaux (1942), Hemirhipus Latreille (1829), Lacais Fleutiaux (1942), Lycoreus Candèze (1857), Mocquerysia Fleutiaux (1899), Neocalais Girard (1971), Pherhimius Fleutiaux (1942), Phibisa Fleutiaux (1942), Propalaus gen. nov., Pseudocalais Girard (1971), Saltamartinus Casari (1996), Tetrigus Candèze (1857) and Thoramus Sharp (1877). The species included in Alaus do not make a monophyletic group and Propalaus gen. nov. is established to include Alaus alicii (Pjatakowa 1941) and A. haroldi (Candèze 1878). A description of Propalaus gen. nov. (type-species: Chalcolepidius haroldi Candèze, 1878) and a new key to Hemirhipini genera are also presented.

Keywords: american species; genera of the world; key to genera; new combinations; type-species.

RESUMO

Esse artigo apresenta uma análise cladística da tribo Hemirhipini. Estão incluídos na análise, 20 gêneros de Hemirhipini (sensu Casari-Chen 1994), Saltamartinus Casari (1996b) (Hemirhipini), 6 gêneros excluídos de Hemirhipini e mantidos em Pyrophorinae (= Agrypninae) (Casari-Chen 1993) e também, Aphileus Candèze (1857), Pyrophorus Billberg (1820) e Thoramus Sharp (1877). As espécies-tipo da maioria dos gêneros e todas as espécies dos gêneros Americanos (exceto Saltamartinus viduus (Chevrolat 1867)) foram analisadas. Essa análise demonstrou que 30 gêneros pertencem a Hemirhipini: Abiphis Fleutiaux (1926), Alaolacon Candèze (1865), Alaomorphus Hauser (1900), Alaus Eschscholtz (1829), Aliteus Candèze (1857), Anthracalaus Fairmaire (1888), Aphileus Candèze (1857), Austrocalais Neboiss (1967), Calais Castelnau (1836), Catelanus Fleutiaux (1942), Chalcolepidius Eschscholtz (1829), Chalcolepis Candèze (1857), Conobajulus Van Zwaluwenburg (1940), Coryleus Fleutiaux (1942), Cryptalaus Ôhira (1967), Eleuphemus Hyslop (1921), Eumoeus Candèze (1874), Fusimorphus Fleutiaux (1942), Hemirhipus Latreille (1829), Lacais Fleutiaux (1942), Lycoreus Candèze (1857), Mocquerysia Fleutiaux (1899), Neocalais Girard (1971), Pherhimius Fleutiaux (1942), Phibisa Fleutiaux (1942), Propalaus gen. nov., Pseudocalais Girard (1971), Saltamartinus Casari (1996), Tetrigus Candèze (1857) e Thoramus Sharp (1877). As espécies incluídas em </>Alaus não formam um grupo monofilético e o gênero Propalaus gen. nov. foi estabelecido para incluir Alaus alicii (Pjatakowa 1941) e A. haroldi (Candèze 1878). A descrição de Propalaus gen. nov. (espécie-tipo: Chalcolepidius haroldi Candèze, 1878) e uma nova chave para os gêneros de Hemirhipini também estão presentes.

Palavras-chave: espécies americanas; espécies-tipo; gêneros do mundo; novas combinações.

INTRODUCTION

The difficulties on Elateridae classification have been pointed out since Lacordaire (1857). Even today, the classification of the higher taxa remains in an unsettled state. The same occurs with the Hemirhipini (Agrypninae) where the number of genera arranged into this tribe is uncertain.

The name "Hémirhipides" was used first by Candèze (1857) to denominate a group formed by Alaus Eschscholtz (1829), Calais Castelnau (1836), Chalcolepis Candèze (1857), Ctenicera Latreille (1829), Euphemus Castelnau (1836), Hemirhipus Eschscholtz (1829), Lycoreus Candèze (1857) and Tetrigus Candèze (1857). He also established Aliteus and Aphileus in "Mélanactides". He put Chalcolepidius Eschscholtz (1829) in "Chalcolépidiides" together with Campsosternus Latreille (1834), Oistus Candèze (1857) and Semiotus Eschscholtz (1829).

Lacordaire (1857) kept the name "Hémirhipides" to include Alaus Eschscholtz, (1829), Ctenicera Latreille (1829), Euphemus Castelnau (1836) and Hemirhipus Eschscholtz (1829). The genera Iphis and Calais, established by Castelnau (1836), were considered as synonyms of Alaus. Chalcolepidius Eschscholtz (1829), Semiotus Eschscholtz (1829) and Campsosternus Latreille (1834) were kept in the "Chalcolépidiides".

The name "Hémirhipides" was changed to "Alaites" by Candèze (1874), that considered Alaus, the most numerous in species and with the widest geographical distribution, as the type of the tribe. He established Eumoeus, considered Calais as a synonym of Alaus and transferred Aliteus Candèze (1857) and Alaolacon Candèze (1865), originally in "Mélanactides", to "Alaites".

Candèze (1891) included Anthracalaus Fairmaire (1888) in "Alaites" and catalogued 11 genera to the tribe: Alaolacon Candèze (1865), Alaus Eschscholtz (1829), Aliteus Candèze (1857) Anthracalaus Fairmaire (1888), Chalcolepis Candèze (1857), Ctenicera Latreille (1829), Eumoeus Candèze (1874), Euphemus Castelnau (1836), Hemirhipus Eschscholtz (1829), Lycoreus Candèze (1857) and Tetrigus Candèze (1857).

Reitter (1905) working on european Coleoptera included Alaus in Hemirhipini and Tetrigus in Ludiini.

Schwarz (1906) followed Candèze (1891), also included Alaomorphus Hauser (1900) in "Alaites" and transferred Alaolacon to the appendix of his work with other "incertae-sedis" genera. He also considered Ludioctenus Fairmaire (1893) as a synonym of Tetrigus.

Heyne & Tachenberg (1908) kept Alaus, Ctenicera and Hemirhipus in Alaini and Chalcolepidius in Chalcolepidiini.

Hyslop (1917) reduced the tribe to subtribal rank, Hemirhipina, that included the Alaites of Candèze (1891), and together with Chalcolepidiina belonged to Pyrophorini. This position was followed by Leng (1920).

Fleutiaux (1919) raised the status of the group to subfamily Hemirhipinae including Alaomorphus, Alaus and Euphemus.

The name Euphemus was preoccupied by a mollusc (Rafinesque 1815) and was replaced by Eleuphemus Hyslop (1921).

Schenkling (1925) catalogued in Hemirhipinae, besides the genera of Schwarz (1906), also Alaolacon and Ludioctenus Fairmaire (1893). He did not consider the synonymization of Schwarz (1906).

Du Buysson (1926) and Fleutiaux (1947) accepted the synonymization of Ludioctenus under Tetrigus.

Fleutiaux (1942) treating of "Les Hemirhipini de la région Malgache" established Coryleus to C. desruisseauxi, Phibisa to two Ctenicera species, Lacais to four Alaus species and Catelanus, Fusimorphus and Pherhimius to Hemirhipus species. He also included in this tribe, Abiphis, Calais, Lycoreus and Mocquerysia. In (1947) assigned Anthracalaus to Ctenicerinae (= Corymbitinae).

Arnett et al. (1969) considered Hemirhipini as synonym of Pyrophorini; Alaus and Chalcolepidius were included in Chalcolepidiini. The tribe Pyrophorini was considered separated from Hemirhipini by Costa (1975).

Laurent (1973) divided the group into subfamilies Hemirhipinae and Alauinae.

Stibick (1979) ressurected the tribes Hemirhipini and Chalcolepidiini.

Casari-Chen (1985) studying the neotropical genera of Hemirhipini transferred Chalcolepidius to this tribe. In (1993), based especially on the presence of subapical tooth at mandibles and separated parameres of aedeagus, removed six genera from the tribe: Alaolacon, Aliteus, Mocquerysia, Eumoeus, Anthracalaus and Tetrigus. According to her, these genera do not make a monophyletic group and they present an uncertain position inside Pyrophorinae (= Agrypninae). She stated that it is quite certain that Alaolacon should be removed from this subfamily. In (1994) she presented a generic phylogenetic analysis to the tribe, including 20 genera: Abiphis, Alaomorphus, Alaus, Austrocalais, Calais, Catelanus, Chalcolepidius, Chalcolepis, Conobajulus, Coryleus, Eleuphemus, Fusimorphus, Hemirhipus, Lacais, Lycoreus, Neocalais, Paracalais, Pherhimius, Phibisa and Pseudocalais. This analysis shows that several genera needed revision and later all american genera were revised (Casari-Chen 1991; Casari 1996a, 1996b, 1998, 1999, 2002a, 2002b) and Saltamartinus Casari (1996) was established.

Calder (1990) considered Aphileus Candèze (1857) more closely related to Hemirhipini genera than pseudomelanactine genera and suggested that Pseudomelanactes Mathieu (1961) is a synonym of Anthracalaus. At the end of this paper he presented "a tentative tribal arrangement of the Australian agrypnine-tetralobine genera" where Aphileus was included in Hemirhipini and Anthracalaus in Pseudomelanactini. In (1992) Calder & Hayek synonymized Pseudomelanactes under Anthracalaus and Pseudomelactini under Hemirhipini. Calder (1996), treating of the genera of Australian Elateridae included Anthracalaus, Aphileus, Austrocalais, Paracalais and Tetrigus in Agrypninae.

Ôhira (1990) considered Paracalais Neboiss (1967) as a synonym of Cryptalaus Ôhira (1967).

Costa (1992) transferred Thoramus Sharp (1877) from Ludiinae to Pyrophorinae (= Agrypninae).

Johnson (2001) stated that "The Hemirhipini includes 28 genera worldwide". In (2002) kept Anthracalaus in Pseudomelanactini and Chalcolepidius in Hemirhipini, both included in Agrypninae.

Casari (2003) described four new species of Alaus and transferred Chalcolepidius alicii Pjatakowa (1941) and C. haroldi Candèze (1878) to this genus. Herein, Propalaus gen. nov., is established to include these two species.

The present analysis demonstrated that Hemirhipini is composed by 30 genera: Abiphis Fleutiaux (1926), Alaolacon Candèze (1865), Alaomorphus Hauser (1900), Alaus Eschscholtz (1829), Aliteus Candèze (1857), Anthracalaus Fairmaire (1888), Aphileus Candèze (1857), Austrocalais Neboiss (1967), Calais Castelnau (1836), Catelanus Fleutiaux (1942), Chalcolepidius Eschscholtz (1829), Chalcolepis Candèze (1857), Conobajulus Van Zwaluwenburg (1940), Coryleus Fleutiaux (1942), Cryptalaus Ôhira (1967), Eleuphemus Hyslop (1921), Eumoeus Candèze (1874), Fusimorphus Fleutiaux (1942), Hemirhipus Latreille (1829), Lacais Fleutiaux (1942), Lycoreus Candèze (1857), Mocquerysia Fleutiaux (1899), Neocalais Girard (1971), Pherhimius Fleutiaux (1942), Phibisa Fleutiaux (1942), Propalaus gen. nov., Pseudocalais Girard (1971), Saltamartinus Casari (1996), Tetrigus Candèze (1857) and Thoramus Sharp (1877).

MATERIAL AND METHODS

The taxa included in this analysis belong to Agrypninae and are listed in ^{Appendix 1}. In this study are included the 20 genera of Hemirhipini, sensu Casari-Chen (1994), 6 genera excluded from this tribe (Casari-Chen,1993), Saltamartinus Casari (1996), Aphileus Candèze (1857), tentatively transferred to Hemirhipini by Calder (1990), Thoramus Sharp (1877), transferred to Pyrophorinae (= Agrypninae) by Costa (1992), and Pyrophorus Bilberg (1820). The type-species of the majority of genera and all species of the American genera (except Saltamartinus viduus (Chevrolat 1867)) are included in the analysis. As out groups are used Alaolacon cyanipennis Candèze (1865), Aliteus reichei (Candèze1857), Anthracalaus westermanni (Candèze 1857), Aphileus lucanoides Candèze (1857), Eumoeus murray Candèze (1874), Mocquerysia coerulipennis Fleutiaux (1929), Pyrophorus diver-gens Eschscholtz (1829), Tetrigus parallelus Candèze (1857) and Thoramus wakefieldi (Sharp 1877).

Cladistic Analysis

One hundred and seven characters were selected, most of which are based on external morphology. Multistate characters were used, always treated as unordered. One hundred twenty two taxa were included in the data matrix representing the Hemirhipini and other Agrypninae genera.

The matrix (^{Appendix 2}) was edited by Nexus program, version 0.5.0 (Page 2001) and the missing data were represented by "?". The trees were conducted using the TNT program (Goloboff et al. 2003), and represented through Winclada version 1.00.08 (Nixon 2002). All characters are treated as unweighted. The trees were rooted a posteriori (Nixon & Carpenter 1993), in Pyrophorus divergens (Agrypninae, Pyrophorini). The analyses were conducted based on 15000 random addition sequences with 2 trees save per replication. The swapping algorithm used was tree bisection reconnection (TBR).

The branches nomenclature for group of species follows Amorim (1982).

Discussion of Characters

1. Body shape (CI 0.20 RI 0.78): (0) wide and arched; (1) narrow, not arched; (2) fusiform. The body in the majority of Hemirhipini is narrowed and not arched; in one group formed by Catelanus trilineatus and Fusimorphus submetallicus is fusiform, representing a synapomorphy shared by species of this group; in Lacais species, three groups of Chalcolepidius species, supremus+, porcatus+ and aurulentus+, C. ferratovittatus, C. serricornis and C. desmaresti, it is wide and arched.

2. Transversal median region of pronotum (CI 0.33 RI 0.00): (0) as wide or slightly narrower than elytral base; (1) strongly narrower than elytral base. The Hemirhipini present the transversal median region of pronotum as wide or slightly narrowed than base of elytra, except Mocquerysia coerulipennis, Alaus latipennis and A. thoracopunctatus, strongly narrower than elytral base.

3. General pubescence (CI 0.37 RI 0.83): (0) simple; (1) scale-like; (2) velvet-like and scale-like; (3) velvet-like and slightly scale-like setae.

Three kinds of setae are present in Hemirhipini: simple, scale-like and velvet-like. Simple setae are thin and piliform; scale-like setae are dorsoventrally flattened, wide and decumbent, with rounded apex and covering the integument color; velvet-like setae are moderately wide, upwardly directed with sharpened apex. One or two kinds of setae are present in each studied species. Simple setae are present in Anthracalaus westermanni, Aphileus lucanoides, Eumoeus murray, Alaolacon cyanipennis, Mocquerysia coerulipennis, Thoramus wakefieldi, Tetrigus parallelus and one group formed by Pherhimius species, Catelanus trilineatus, Fusimorphus submetallicus and Saltamartinus species. Scale-like setae are present in the majority of the more derived groups of the Hemirhipini; among the more basal, they are present only in Eleuphemus species. Velvet-like and scale-like setae are present in Abiphis nobilis, Lycoreus goudoti, Pseudocalais basilewskyi, one group of Alaus species, myops+ and Alaus tricolor; velvet-like and slightly scale-like setae represents one synapomorphy shared by Hemirhipus species.

4. Thickness of scale-like pubescence (CI 0.40 RI 0.75): (0) fine; (1) moderate ("normal"); (2) strong (wide).

The scale-like seta is considered fine when it is only slightly wider than simple setae; moderate when it is moderately wider than simple setae, and strong when it is very wider than simple setae. Scale-like setae fine are present in Hemirhipus species, Phibisa pupieri and Coryleus pectinatus. The majority of Hemirhipini presents scale-like pubescence with moderate thickness, considered as "normal" and representing a synapomorphy shared by group Aliteus reichei+. Scale-like setae wide are present in Eleuphemus species and Alaus candezei.

5. Pubescence of male pronotum(CI 0.33 RI 0.67): (0) forming longitudinal stripes; (1) forming eye-like spots; (2) forming longitudinal elliptical bands near margins; (3) forming median basal spot; (4) unicolor; (5) forming triangular area near hind angles; (6) forming longitudinal lateral bands not reaching lateral margin; (7) forming irregular small spots; (8) forming basal spots; (9) forming longitudinal median elliptical spot. The pubescence coloration of the male pronotum is unicolor or forms several patterns in different groups. In three groups of Chalcolepidius species, albiventris+, copulatuvittatus+ and approximatus+ (except angustus+), C. tartarus and C. virginalis the pubescence forms longitudinal stripes; in Eleuphemus species, Hemirhipus ochraceipilosus, Cryptalaus prosectus, one group formed by Neocalais macer and Austrocalais pogonodes, Propalaus haroldi and one group of Alaus species, cinnamomeus+, forms eye-like spots; in Hemirhipus species (except H. ochraceipilosus), one group of Chalcolepidius species, erythroloma+, C. albisetosus and C. villei forms longitudinal elliptical band near margins; in one group of Chalcolepidius species, angustatus+, C. serricornis and C. trucuvittatus forms median basal spot; in Anthracalaus westermanni, Aphileus lucanoides, Eumoeus murray, Alaolacon cyanipennis, Mocquerysia coerulipennis, Thoramus wakefieldi, Tetrigus parallelus, one group formed by Pherhimius species, Catelanus trilineatus, Fusimorphus submetallicus and Saltamartinus it is unicolor; in Chalcolepidius validus and C. virgatipennis, forms triangular area near hind angles; in Lacais glauca, longitudinal lateral bands not reaching lateral margins; in Aliteus reichei and Lacais nietoi, irregular small spots; in Alaomorphus candezei, basal spots; in Abiphis nobilis, Lycoreus goudoti and Pseudocalais basilewskyi, longitudinal median spot.

6. Velvet-like pubescence on pronotum (CI 1.0 RI 1.0): (0) forming two longitudinal elliptical spots; (1) forming two eye-like spots; (2) forming one median elliptical spot.

The velvet-like pubescence of the pronotum forms two longitudinal elliptical spots, of varied sizes, in Hemirhipus species; in Pseudocalais basilewskyi, one group of Alaus species, myops+ and A. tricolor, two eye-like spots; in Abiphis nobilis and Lycoreus goudoti, one longitudinal median elliptical spot.

7. Two longitudinal elliptical velvet-like spots on pronotum (CI 0.75 RI 0.0): (0) almost as wide as half of pronotum width and reaching both extremities; (1) slightly narrower than half of pronotum width and reaching base; (2) very narrower and shorter than half of pronotum; (3) very small (about 1/5 pronotum length).

The longitudinal elliptical velvet-like spots on pronotum are the different sizes and shapes and are located in different positions, depending on the species. They are almost as wide as half of pronotum width and reaching both extremities in Hemirhipus rojasi and H. fairmairii; slightly narrower than half of pronotum width and reaching base in Hemirhipus lineatus, representing an autapomorphy to this species; very narrower and shorter than half of pronotum in one group of Hemirhipus species, bimaculatus+, representing a synapomorphy shared by species of this group; very small, about 1/5 pronotum length in Hemirhipus ochraceipilosus, representing an autapomorphy to this species.

8. Pubescence of eye-like spots on pronotum (CI 0.33 RI 0.66): (0) scale-like setae; (1) velvet-like setae. The pubescence of the eye-like spots on pronotum is clothed by different kinds of setae: scale-like in Eleuphemus species, Cryptalaus prosectus, one group formed by Neocalais macer and Austrocalais pogonodes, Propalaus haroldi, three groups of Alaus species, cinnamomeus+, latipennis+ and calcaripiosus+, A. nobilis and A. thoracopunctatus; velvet-like in Pseudocalais basilewskyi, one group of Alaus species, myops+ and A. tricolor.

9. Eye-like spots on pronotum (CI 0.75 RI 0.75): (0) without whitish contour; (1) surrounded by narrow whitish band; (2) included in the white lateral band; (3) inside elliptical spots.

Usually the eye-like spots on pronotum are not surrounded by whitish bands or spots. In one group of Alaus species, myops+ and A. nobilis, they are surrounded by narrow whitish band; in Alaus zunianus they are included in white lateral bands; in Alaus patricius they are inside elliptical spots.

10. White pubescence of elytra (CI 0.25 RI 0.52): (0) continuous or forming uneven spots; (1) forming small patches; (2) white totally; (3) forming longitudinal bands; (4) forming longitudinal bands; (5) forming longitudinal lateral bands.

The white pubescence of elytra, are continuous covering the elytra totally or forms different patterns. It is continuous or forming uneven patterns in Eleuphemus fasciatus, Alaomorphus candezei, Lycoreus goudoti, Alaus latipennis, A. myops, A. lusciosus, one group of Chalcolepidius species, albisetosus+ and C. virginalis, C. sulcatus, C. validus, C. erythroloma and C. rugatus; forming small patches in one group of Alaus species, calcaripilosus+, representing a synapomorphy shared by species of this group; white totally in Propalaus haroldi, representing an autapomorphy to this species; forming longitudinal bands in Alaus unicus, some Chalcolepidius species from groups supremus+, aurulentus+ and C. tartarus; forming longitudinal lateral bands in the majority of Chalcolepidius species from group fasciatus+, C. albiventris, C. truncuvittatus and C. porcatus.

11. Pubescence coloration of male and female (CI 0.33 RI 0.33): (0) similar; (1) different.

The pubescence in the majority of Hemirhipini is similar in male and female. Lacais nietoi, L. glauca and one group of Chalcolepidius species, mexicanus+, present the coloration of male pubescence different from that of female.

12. Longitudinal lateral stripes band-like of pubescence on pronotum (CI 0.40 RI 0.40): (0) wide and reaching hind angles; (1) moderately wide and reaching hind angles; (2) narrow and reaching hind angles; (3) fused at base; (4) narrowed at extremities.

In several Hemirhipini species the pubescence of pronotum forms one longitudinal lateral band each side of pronotum. These bands varies in size and shape. They are wide and reach hind angles of pronotum in Alaus myops, A. zunianus and Chalcolepidius corpulentus; moderately wide and reaching hind angles in Alaus oculatus, two groups of Chalcolepidius species, albiventris+ and approximatus+ and C. tartarus; narrow and reaching hind angles in Lacais glauca, Alaus oculatus, Chalcolepidius truncuvittatus, C. spinipennis, C. virginalis, C. virgatipennis and C. fasciatus; fused at base, representing an autopomorphy to Chalcolepidius copulatuvittatus; narrowed at extremities representing a synapomorphy shared by one group of Chalcolepidius species, angustatus+.

13. Coloration of elytral pubescence (CI 0.25 RI 0.57): (0) unicolor; (1) striped with or without lateral bands; (2) with lateral bands; (3) with spots; (4) with sinuous transverse bands; (5) apical region of different color; (6) with small patches; (7) with bands and spots; (8) with longitudinal elliptical and transverse bands; (9) with longitudinal and transverse bands.

The coloration of elytral pubescence forms characteristic patterns in several Hemirhipini groups. It is unicolor in Anthracalaus westermanni, Aphileus lucanoides, Eumoeus murray, Alaolacon cyanipennis, Mocquerysia coerulipennis, Thoramus wakefieldi, Tetrigus parallelus, one group formed by Catelanus trilineatus and Fusimorphus submetallicus, Hemirhipus ochraceipilosus, H. rojasi, H. bimaculatus, Conobajulus ugiensis, Lacais suturalis, Phibisa pupieri, Coryleus pectinatus, Propalaus and Chalcolepis species, Alaus veracruzanus, A. cinnamomeus, A. sericeus, A. nobilis, five groups of Chalcolepidius species, mexicanus+, dugesi+, desmaresti+, viridipilis+ and lacordairii+, C. supremus, C. oxydatus and C. rubripennis; striped with or without lateral bands in one group of Hemirhipus species, fairmairii+, Alaus unicus, three groups of Chalcolepidius species, extenuatuvittatus+, albisetosus+ and aurulentus+ (except erythroloma+, C. corpuletus and apacheanus+) and C. virginalis, C. porcatus and C. validus; with lateral bands in Neocalais macer, two groups of Chalcolepidius species, erythroloma+ and apachenus+, and some Chalcolepidius species of other different groups; with spots in Lacais nietoi, Alaus tricolor and A. myops; with sinuous transverse bands in Pherhimius species, Pseudocalais basilewskyi and Alaus patricius; with apical region of different color in one group of Hemirhipus species, apicalis+ and Alaomorphus candezei; with small patches in Aliteus reichei, one group formed by Calais excavatus and Cryptalaus prosectus, two groups of Alaus species, calcaripilosus+ and melanops+ and A. latipennis; with bands and spots in Eleuphemus species, Austrocalais pogonodes, Lacais glauca, Lycoreus goudoti and Alaus thoracopunctatus; with longitudinal elliptical and transverse band representing one synapomorphy shared by Saltamartinus species; with longitudinal and transverse bands representing one autapomorphy to Abiphis nobilis.

14. Epipleura pubescence coloration (CI 0.26 RI 0.38): (0) similar to underside; (1) different from underside; (2) different from underside, pronotum with lateral stripes; (3) different from underside, pronotum and elytra with lateral stripes; (4) partially different from underside. The pubescence coloration of epipleura is usually similar to underside.

It is different from underside and pronotum without lateral stripes in Lacais species, one group of Alaus species, cinnamomeus+, A. tricolor and Chalcolepidius aurulentus; different from underside and pronotum with lateral stripes in Chalcolepidius copulatuvittatus, C. validus, C. approximatus and C. rostainei; different from underside, pronotum and elytra with lateral stripes in one group of Chalcolepidius species, limbatus+, C. tartarus and C. corpulentus; partially different from underside, pronotum without stripes in one group of Chalcolepidius species, extenuatuvittatus+ and C. virgatipennis.

15. Pubescence of ventrites (CI 0.25 RI 0.00): (0) unicolor; (1) with white patches.

Only the ventrites of Alaomorphus candezei, Chalcolepidius tartarus, C. approximatus and C. apacheanus present white patches, especially near lateral margins.

16. Coloration of pronotal integument (CI 0.62 RI 0.70): (0) unicolor; (1) forming three longitudinal bands; (2) forming two elliptical or rounded spots; (3) with longitudinal median band; (4) forming eye-like spots; (5) forming one median elliptical spot.

The integument of pronotum is usually unicolor; it forms three longitudinal bands in Pherhimius dejeani and Catelanus trilineatus; two elliptical or rounded spots in Eleuphemus species, one group of Hemirhipus species, bimaculatus+ and Conobajulus ugiensis; longitudinal median band, representing an autapomorphy to Propalaus alicii; eye-like spots, representing an autapomorphy to Alaus calacaripilosus; one median elliptical spot, representing an autapomorphy to Abiphis nobilis.

17. Frons (CI 0.36 RI 0.50): (0) not carinate; (1) carinate; (2) slightly carinate; (3) incompletelly carinate; (4) not carinate and forming a rounded fold.

The frons is usually not carinate: the ridge between frons and nasal are absent. In Thoramus wakefieldi, one group formed by Pherhimius, Catelanus, Fusimorphus and Saltamartinus species, it is carinate and the ridge is present. In Eleuphemus species, Hemirhipus bimaculatus and H. fairmairii, it is slightly carinate, forming a very weak ridge; in Anthracalaus westermanni, Tetrigus parallelus, Hemirhipus hougeti and Pseudocalais basilewskyi, it is incompletely carinate, and the ridge is absent at middle of frons. In Phibisa pupieri the frons is not carinate but the anterior margin is folded, representing an autapomorphy to this species.

18. Fore angles of frons (CI 0.28 RI 0.68): (0) normal; (1) raised; (2) slightly prominent. In the majority of the studied species, the fore angles of frons are almost horizontal, not raised, considered as normal. In Saltamartinus perroudi, Austrocalais pogonodes, Chalcolepis species, one group of Alaus species, latipennis+, A. unicus and A. calcaripilosus, the fore angles are raised; it is slightly prominent in Hemirhipus species, representing a synapomorphy shared by species of this genus.

19. Basal region of frons (CI 0.50 RI 0.88): (0) smooth; (1) with median tubercle. In the majority of studied species the frons is smooth; Saltamartinus decorus and Hemirhipus species present a small tubercle at median basal region of frons.

20. Nasal (CI 0.23 RI 0.75): (0) very short; (1) high; (2) absent; (3) sloped.

The nasal is absent when the frons is at same level that the base of labrum; representing a synapomorphy shared by all Hemirhipini. It is very short in one and more derived group, formed by Propalaus, Alaus and Chalcolepidius species; it is high in a group formed by Thoramus wakefield, Tetrigus parallelus, and Pherhimius, Catelanus, Fusimorphus, Saltamartinus and Hemirhipus and Lacais species, Pseudocalais basilewskyi, Propalalus alicii, Alaus sericeus, A. nobilis, A. calcaripilosus and A. zunianus. In two groups, it is inclined from dorsal region of frons to basal region of labrum, considered as sloped: one group formed by Calais excavatus and Cryptalaus prosectus and other, by Alaomorphus candezei, Abiphis nobilis, Phibisa pupieri and Coryleus pectinatus.

21. Number of antennomeres of male (CI 0.20 RI 0.83): (0) 11; (1) 12.

The antennae of male in the majority of studied species have 11 antennomeres. In three groups, the antennae have 12 antennomeres: Eumoeus murray, Mocquerysia coerulipennis, Eleuphemus fasciatus and E. funerarius; Pherhimius, Catelanus, Fusimorphus, Saltamartinus and Hemirhipus species; and Lacais species (except L. suturalis), Alaomorphus candezei, Abiphis nobilis and Phibisa pupieri.

22. Antennal shape of male (CI 0.22 RI 0.74): (0) serrate; (1) strongly serrate; (2) pectinate; (3) flabellate; (4) biflabellate.

The antennae are usually serrate. They are considered strongly serrate when the antennomeres are prominent lateroexternally representing an intermediary state between pectinate and serrate. Antennae strongly serrate are present in Anthracacalus westermanni, Propalaus haroldi, one group of Alaus species, sericeus+, two groups of Chalcolepidius species, supremus+ and rubripennis+ (except porcatus+); pectinate in Propalaus alicii, Chalcolepidius mexicanus, C. dugesi and C. inops; flabellate in one group formed by Tetrigus parallelus, and Pherhimius, Catelanus, Fusimorphus, Saltamartinus and Hemirhipus species, one group formed by Lacais species (except L. suturalis), Alaomorphus candezei, Abiphis nobilis, Phibisa pupieri, Coryleus pectinatus and Lycoreus goudoti, Alaus patricius, Chalcolepidius viridipennis and C. smaragdinus; biflabellate representing a synapomorphy shared by group formed by Eumoeus murray, Alaolacon cyanipennis, Mocquerysia coerulipennis, Eleuphemus species.

23. Antennae (CI 0.60 RI 0.70): (0) serrate, pectinate or flabellate from 3rd antennomere; (1) serrate, pectinate or flabellate from 4th antennomere.

The antennae shapes, serrate, pectinate or flabellate, start at 3^rd or 4^th antennomere. They start at 3^rd antennomere in two groups, one formed by Eumoeus murray, Alaolacon cyanipennis, Mocquerysia coerulipennis and Eleuphemus species, and other by Pherhimius, Catelanus, Fusimorphus and Saltamartinus species, Aliteus reichei, Conobajulus ugiensis, Calais excavatus, one group formed by Neocalais macer and Austrocalais pogonodes, one group formed by Pseudocalais basilewskyi, Propalaus, Chalcolepis and Alaus (except group sericeus+) and many Chalcolepidius species, especially from group desmaresti+. They start at 4^th antennomere in Anthracalaus westermanni, Aphileus lucanoides, Thoramus wakefieldi, Tetrigus parallelus, Hemirhipus species, Crypatalaus prosectus, one group formed by Lacais species, Alaomorphus candezei, Abiphis nobilis, Phibisa pupieri, Coryleus pectinatus and Lycoreus goudoti, Propalaus species, one group of Alaus species, sericeus+, two groups of Chalcolepidius species, ferratuvittatus+ and viridipilis+, C. albiventris, C. virginalis and C. inops.

24. 3rd antennomere of male (CI 0.29 RI 0.72): (0) triangular-elongate; (1) triangular-short; (2) transverse; (3) transverse with spiniform appendix; (4) flabellate; (5) transverse, prominent laterally; (6) biflabellate; (7) elongate.

The 3^rd antennomere is variable in shape and is characteristic in several groups. It is elongate and triangular in the majority of species, especially those of Alaus and Chalcolepidius; it is short and triangular in Anthracalaus westermanni, Calais excavatus, Propalalus haroldi, one group of Chalcolepis species, similis+, Alaus unicus, two groups of Chalcolepidius species, supremus+ and viridipilis+, C. rubripennis and C. virginalis; wider than long, designated as transverse, in Thoramus wakefieldi, Tetrigus parallelus and some species of Chalcolepidius; transverse, prominent laterally forming an spiniform appendix in a group formed by Lacais species, Alaomorphus candezei, Abiphis nobilis, Phibisa pupieri, Coryleus pectinatus and Lycoreus goudoti, Propalalus alicii, Alaus nobilis and A. patricius; flabellate, representing a synapomorphy shared by group formed by and Pherhimius, Catelanus, Fusimorphus, Saltamartinus and Hemirhipus species; transverse and prominent laterally in Cryptalaus prosectus, one group of Alaus species, sericeus+; biflabellate, representing a synapomorphy shared by group formed by Eumoeus murray, Alaolacon cyanipennis, Mocquerysia coerulipennis, Eleuphemus fasciatus and E. funerarius; elongate, representing an autapomorphy to Aphileus lucanoides.

25. 3^rd antennomere (CI 0.15 RI 0.67): (0) shorter than 4th; (1) longer than 4th; (2) as long as 4th.

Usually the 3^rd antennomere is shorter than 4^th; it is longer than 4^th in Aphileus lucanoides, one group formed by Alaolacon cyanipennis and Mocquerysia coerulipennis, Catelanus trilineatus, Aliteus reichei, Chalcolepis austerus, one group of Chalcolepidius species, lafargi+, C. lacordairii and C. fabricii; 3^rd as long as 4^th in Eumoeus murray, Eleuphemus species, two groups, one formed by Pherhimius, Catelanus, Fusimorphus, Saltamartinus and Hemirhipus species, and other by Lacais species, Alaomorphus candezei, Abiphis nobilis and Phibisa pupieri.

26. Mandibles (CI 1.00 RI 1.00): (0) very long, prominent; (1) short, not promient.

Usually the mandibles are short and not prominent; in Aphileus lucanoides they are very long and prominent, representing an autapomorphy to this species.

27. Subapical tooth of mandible (CI 0.33 RI 0.71): (0) present; (1) absent.

Usually the subapical tooth of mandibles is absent; it is present in the basal groups: Anthracalaus westermanni, Aphileus lucanoides, Eumoeus murray, Alaolacon cyanipennis, Mocquerysia coerulipennis, Thoramus wakefieldi and Aliteus reichei.

The presence of subapical tooth of mandibles was used by Casari-Chen (1993, 1994) to remove some genera from Hemirhipini. The absence of subapical tooth of mandibles was considered a synapomorphy to the tribe.

28. Length of thorax (CI 1.00 RI 1.00): (0) normal; (1) long in relation to abdomen.

The thorax is long in relation to abdomen length only in Lacais species, representing a synapomorphy shared by species of this genus.

29. Luminescent vesicles of pronotum (CI 1.00 RI 1.00): (0) present; (1) absent.

In Hemirhipini the luminescent vesicles are absent, representing a synapomorphy shared by all members of this tribe.

30. Medioanterior margin of pronotum (CI 0.40 RI 0.85): (0) straight or slightly sinuous; (1) raised forming two teeth; (2) sinuous. The anterior margin of pronotum is usually straight or slightly sinuous. It is raised forming two teeth near middle in three groups: one formed by Calais excavatus and Crypatalus prosectus, the second by Pseudocalais basilewskyi, Propalaus species and Chalcolepis species and the third by Alaus group nobilis+ and Alaus thoracopunctatus; it is sinuous in Hemirhipus species, representing a synapomorphy shared by species of this genus.

31. Lateral margins of pronotum (CI 0.15 RI 0.66): (0) slightly sinuous; (1) sinuous; (2) rounded; (3) almost straight; (4) straight.

The majority of studied species, especially Chalcolepidius species, presents lateral margins of pronotum slightly sinuous. They are sinuous in Anthracalaus westermanni, Alaus tricolor and Chalcolepidius rubripennis; rounded in Tetrigus parallelus, Calais excavatus, Neocalais macer, Alaomorphus candezei, one group formed by Coryleus pectinatus and Lycoreus goudoti, some species of Alaus and one group of Chalcolepidius species, dugesi+; almost straight in Mocquerysia coerulipennis, Eleuphemus species, one group formed by Thoramus wakefieldi and Pherhimius, Catelanus, Fusimorphus, Saltamartinus and Hemirhipus species, Conobajulus ugiensis, one group formed by Pseudocalais basilewskyi, Propalaus species, Chalcolepis species and one group of Alaus species, cinnamomeus+.

32. Lateral margins of pronotum (CI 1.00 RI 1.00): (0) raised forming a ridge; (1) not raised.

The lateral margins of pronotum are usually not raised; in Chalcolepis species they are raised in a narrow band forming like a brim, representing a synapomorphy shared by species of this genus.

33. Hind angles of pronotum (CI 0.15 RI 0.52): (0) backwardly or slightly divergent; (1) divergent; (2) strongly divergent.

The hind angles of pronotum are usually backwardly or slightly divergent. They are divergent in Anthracalus westermanni, Aliteus reichei, one group formed by Neocalais macer, Austrocalais pogonodes, Lacais species, Alaomorphus candezei and Phibisa pupieri, Alaus veracruzanus and two groups of Alaus species, sericeus+ and calcaripilosus+; and strongly divergent in Aphileus lucanoides, Mocquerysia coerulipennis, Chalcolepis species, Alaus tricolor and A. thoracopunctatus.

34. Hind angles of pronotum (CI 0.27 RI 0.85): (0) unicarinate; (1) not carinate; (2) carina bifurcate; (3) raised but not forming a carina.

The hind angles of pronotum of the majority of Hemirhipini are not carinate. The basal groups, Anthracalaus westermanni, Aphileus lucanoides, Eumoeus murray, Alaolacon cyanipennis, one group formed by Mocquerysia coerulipennis, Thoramus wakefieldi, Tetrigus parallelus, Pherhimius dejeani, Fusimorphus submetallicus, Saltamartinus species, and Hemirhipus species, Aliteus reichei, one group formed by Conobajulus ugiensis, Calais excavatus and Cryptalaus prosectus, one group formed by Neocalais macer, Austrocalais pogonodes, Alaomorphus candezei, Abiphis nobilis, Phibisa pupieri, Coryleus pectinatus and Lycoreus goudoti, Alaus patricius, one group of Alaus species, calcaripilosus+, one group of Chalcolepidius species, dugesi+, present one carina at middle; Pherhimius fascicularis presents carina bifurcate, representing an autapomorphy to this species; hind angles raised but not forming a ridge in Eleuphemus species, Propalaus haroldi, Chalcolepis and Alaus species.

35. Apex of hind angles of pronotum (CI 1.00 RI 1.00): (0) normal; (1) constricted.

Only Fusimorphus submetallicus presents constricted apex of hind angles of pronotum, representing an autapomorphy to this species; in the remaining Hemirhipini the apex of hind angles of pronotum is slightly narrowed apicad.

36. Convexity of pronotum (CI 0.17 RI 0.44): (0) slight or moderate; (1) stronger at middle; (2) stronger medioanteriorly; (3) forming longitudinal median ridge; (4) strongly convex from sutures; (5) moderately convex from sutures. The pronotum of the majority of Hemirhipini is slightly or moderately convex from sutures.

In some groups, the convexity varies in intensity and in location. It is stronger convex at middle, but not forming a longitudinal median ridge in Austrocalalis pogonodes, Propalaus haroldi, Chalcolepis species (except C. similis), Alaus latipennis, two groups of Chalcolepidius species, dugesi+ and boucardi+, C. copulatuvittatus and C. rubripennis; stronger convex medioanteriorly and the convexity decreases basad in Pseudocalais basilewskyi and Chalcolepidius inops; stronger convex at middle forming a longitudinal median ridge in Neocalais macer, Abiphis nobilis, Phibisa pupieri, Propalaus alicii, Chalcolepis similis and Chalcolepidius desmaresti; strongly convex from sutures in one group formed by Eumoeus murray, Alaolacon cyanipennis and Eleuphemus species, Tetrigus parallelus, Fusimorphus submetallicus, Hemirhipus species, Cryptalaus prosectus, Alaomorphus candezei, Coryleus pectinatus and some Alaus species; moderately convex from sutures in Mocquerysia coerulipennis and Aliteus reichei.

37. Pronotum (CI 0.60 RI 0.83): (0) without furrow; (1) slightly grooved longitudinal medially; (2) with two longitudinal grooves; (3) grooved at middle near base

The pronotum of Hemirhipini is usually slight-, moderate- or strongly convex from the sutures, but in some groups the pronotum presents longitudinal grooves near middle. The pronotum is slightly grooved longitudinal medially in one group formed by Eumoeus murray, Alaolacon cyanipennis Mocquerysia coerulipennis and Eleuphemus species and one group of Alaus species, melanops+; with two longitudinal grooves near middle in Neocalais macer and one group formed by Abiphis nobilis, Phibisa pupieri, Coryleus pectinatus and Lycoreus goudoti; grooved at middle near base in Calais excavatus, representing an autapomorphy to this species.

38. Pronotum (CI 1.00 RI 1.00): (0) without carinae; (1) bearing median tranverse carina near base.

In Neocalais macer the pronotum presents a short transverse carina near base, representing an autapomorphy to this species.

39. Median basal tubercle of pronotum (CI 0.21 RI 0.77): (0) transverse; (1) transverse with carina; (2) elongate; (3) triangular elongate or indistinct; (4) raised and rounded.

The majority of Hemirhipini, including Aphileus lucanoides, one group formed by Alaolacon cyanipennis and Mocquerysia coerulipennis, Tetrigus parallelus, Conobajulus ugiensis, one group of Alaus species, melanops+, and one group of Chalcolepidius species, desmaresti+, have median basal tubercle of pronotum triangular elongate or indistinct. In Anthracalaus westermanni, Eumoeus murray, Eleuphemus species, Thoramus wakefieldi, Catelanus trilineatus, Fusimorphus submetallicus, Aliteus reichei, Cryptalaus prosectus, Neocalais macer, Lacais species, Lycoreus goudoti, Chalcolepis species and Alaus species (except A. tricolor, and group melanops+), the median basal tubercle of pronotum is transverse; in one group of Chalcolepidius species, dugesi+, it is transverse with transverse carina; in a group formed by Pherhimius, Saltamartinus and Hemirhipus species, Calais excavatus, Austrocalais pogonodes, one group formed by Alaomorphus candezei, Abiphis nobilis, Phibisa pupieri and Coryleus pectinatus, one group formed by Pseudocalais basilewskyi and Propalaus species, Alaus tricolor and one group of Chalcolepidius species, supremus+, it is elongate.

40. Posterior margin of pronotum (CI 0.33 RI 0.89): (0) rounded at middle; (1) prominent and notched at middle; (2) prominent and straight at middle; (3) notched at middle.

In all Chalcolepis and Chalcolepidius species the posterior margin of pronotum is prominent and notched at middle. In the remaining Hemirhipini, the posterior margin of pronotum is not prominent and notched at middle, except, Aphileus lucanoides, Alaomorphus candezei and Propalaus haroldi, with posterior region prominent and straight at middle, and one group formed by Catelanus trilineatus and Fusimorphus submetallicus, Lacais species, Lycoreus goudoti and Propalaus alicii, posterior region rounded at middle.

41. Prosternum (CI 0.15 RI 0.73): (0) slightly convex from sutures; (1) moderately convex from sutures; (2) strongly convex from sutures; (3) strongly convex, flat longitudinally and grooved laterally.

The convexity of prosternum is variable and starts at sutures or prosternum is slight groove laterally and convexity starts more internally. Prosternum slightly convex and convexity starting from sutures is present in Aphileus lucanoides, Thoramus wakefieldi, Tetrigus parallelus, one group formed by Pherhimius, Catelanus, Fusimorphus, and Saltamartinus species, Lacais species, Pseudocalais basilewskyi, one group of Alaus species, calcaripilosus+, and one group of Chalcolepidius species, supremus+. Prosternum moderately convex starting from sutures is present in Mocquerysia coerulipennis and some Alaus species; strongly convex starting from sutures, in Anthracalaus westermanni, Eumoeus murray, Hemirhipus species, Aliteus reichei, one group formed by Conobajulus ugiensis, Calais excavatus and Cryptalaus prosectus, one group formed by Neocalais macer and Austrocalais pogonodes, Alaomorphus candezei and some Alaus species; strongly convex, flat longitudinally and grooved laterally in a group formed by Alaolacon cyanipennis and Eleuphemus species, one group formed by Phibisa pupieri, Coryleus pectinatus and Lycoreus goudoti, one group formed by Propalaus and Chalcolepis species, one group of Alaus species, nobilis+, and one large group of Chalcolepidius species, desmaresti+.

42. Prosternum (CI 1.00 RI 1.00): (0) gradually widened anteriad; (1) widened frontally; (2) widened at median region.

The prosternum is usually slightly and gradually widened anteriad. In Pseudocalais basilewskyi it is strongly widened frontally and in Alaomorphus candezei, strongly widened at median region, representing an autapomorphy to each species.

43. Notosternal suture (CI 0.26 RI 0.85): (0) straight; (1) slightly sinuous; (2) moderately to strongly sinuous; (3) straight and slightly curved near apex; (4) semicircular.

The notosternal suture is straight in Aphileus lucanoides, Thoramus wakefieldi, one group formed by Pherhimius, Catelanus, Fusimorphus and Saltamartinus species, Aliteus reichei, one group formed by Conobajulus ugiensis, Calais excavatus and Cryptalaus prosectus, one group formed by Neocalais macer, Lacais species, Abiphis nobilis, Phibisa pupieri, Coryleus pectinatus and Lycoreus goudoti, one group formed by Propalaus and Chalcolepis species and Alaus species (except A. nobilis, A. tricolor and A. zunianus); slightly sinuous in Anthracalaus westermanni, one group formed by Alaolacon cyanipennis and Mocquerysia coerulipennis, Tetrigus parallelus, one group of Alaus species, nobilis+, A. zunianus, one group of Chalcolepidius species, supremus+, C. jansoni, C. rubripennis, C. viridipilis, C. chalcantheus and C. boucardi; moderately to strongly sinuous in Eumoeus murray, Eleuphemus species, Austrocalais pogonoides, Pseudocalais basilewskyi, one group of Chalcolepidius species, virginalis+, C. desmaresti, C. tartarus and C. smaragdinus; straight or slightly curved near apex in Hemirhipus species, representing a synapomorphy shared by species of this genus; semicircular in Alaomorphus candezei, representing an autapomorphy to this species.

44. Prosternal channel (CI 0.09 RI 0.81): (0) absent; (1) present.

The notosternal sutures are, in several Hemirhipini groups, opened frontally forming a channel. Prosternal channel is present in Anthracalaus westermanni, Mocquerysia coerulipennis, Eleuphemus species, Aliteus reichei, one group formed by Calais excavatus and Cryptalaus prosectus, Neocalais macer, one group formed by Abiphis nobilis, Phibisa pupieri, Coryleus pectinatus, Lycoreus goudoti, Propalaus species, Alaus veracruzanus, A. latipennis and one large group of Chalcolepidius species, rubripennis+. In the remaining Hemirhini the prosternal channel is absent.

45. Hypomera (CI 1.00 RI 1.00): (0) "normal"; (1) grooved near sutures; (2) slightly grooved near sutures.

The Hemirhipini hypomera are punctuated and slightly concave in whole surface, considered as "normal". In Eleuphemus species, they are from moderate- to strongly grooved near notosternal sutures, representing a synapomorphy shared by species of this genus; in Phibisa pupieri, slightly grooved near notosternal sutures, representing an autapomorphy to this species.

46. Apex of prosternal spine (CI 1.00 RI 1.00): (0) with tooth; (1) rounded.

In all studied Hemirhipini the prosternal spine presents rounded apex, representing a synapomorphy shared by all groups of this tribe. This state had already been used as a synapomorphy for the tribe by Casari-Chen (1994).

47. Basal region of prosternal spine (CI 1.00 RI 1.00): (0) flat or slightly grooved; (1) bearing longitudinal elongate groove.

The basal region of prosternal spine of Hemirhipini is flat or slightly grooved between procoxae; Alaomorphus candezei has a longitudinal elongate groove in this area, representing an autapomorphy to this species.

48. Borders of mesosternal cavity (CI 0.22 RI 0.46): (0) horizontal; (1) declivous; (2) elevate at base.

The borders of mesosternal cavity of the majority of Hemirhipini are horizontal; in Aphileus lucanoides, Catelanus trilineatus, one group formed by Calais excavatus and Cryptalaus prosectus, Lacais and Chalcolepis species, Alaus nobilis, A. latipennis and A. plebejus are declivous; in Conobajulus ugiensis are elevate at base, representing an autapomorphy to this species.

49. Borders of mesosternal cavity horizontal (CI 0.27 RI 0.62): (0) horizontal in whole length; (1) excavate at middle; (2) horizontal and slightly declivous frontally; (3) horizontal and moderately declivous frontally; (4) horizontal and strongly declivous frontally; (5) horizontal with thick margins at base.

Among the Hemirhipini with borders of mesosternal cavity horizontal, in Pherhimius species, the borders are horizontal in whole length, representing a synapomorphy shared by species of this genus; in Anthracalaus westermanni, Eumoeus murray, Tetrigus parallelus, Fusimorphus submetallicus, Saltamartinus species, Aliteus reichei, one group formed by Neocalais macer and Austrocalais pogonodes, Alaomorphus candezei and Alaus calcaripilosus are excavate at middle; in Alaolacon cyanipennis, Eleuphemus species, Thoramus wakefieldi, one group formed by Abiphis nobilis, Phibisa pupieri, Coryleus pectinatus and Lycoreus goudoti and some species of Alaus and some of Chalcolepidius are horizontal and slightly declivous frontally; in Pseudocalais basilewskyi and the majority of Chalcolepidius species are horizontal and moderately declivous frontally; in Mocquerysia coerulipennis, Propalaus species, Alaus cinnamomeus, A. tricolor and one group Alaus species, lusciosus+, are horizontal and strongly declivous frontally; in Hemirhipus species are horizontal with margins thickend at base, representing a synapomorphy shared by species of this genus.

50. Meso-mestasternal suture (CI 0.50 RI 0.98): (0) present; (1) absent or obsolete.

In the majority of Hemirhipini the mesosternum is separated from metasternum by a suture behind the mesosternal cavity. This suture is absent or obsolete in Phibisa pupieri and Chalcolepidius species.

Meso-metastenal suture absent or obsolete was used by Casari (2002b) as a synapomorphy shared by Chalcolepidius species.

51. Metasternal median suture (CI 0.66 RI 0.00): (0) "normal", without furrow or carina; (1) with transverse anterior carina; (2) furrowed near base; (3) forming an elliptical anterior cavity.

In the majority of Hemirhipini the metasternal median suture is straight, with the same width in whole its length, considered as "normal". In Calais excavatus it presents a short tranverse carina anteriorly, behind meso-metasternal suture, representing an autapomorphy to this species; in Cryptalaus prosectus and Austrocalais pogonodes it is furrowed near base; in Pherhimius fascicularis it is furrowed at base or it forms an elliptical anterior cavity.

52. Median anterior region of metasternum (CI 1.00 RI 1.00): (0) flat; (1) forming a prominence between mesocoxae.

In the Hemirhipini the median anterior region of metasternum, near mesocoxae, is flat; only in Conobajulus ugiensis this area is raised forming a rounded prominence, representing an autapomorphy to this species.

53. Metacoxal plate (CI 0.33 RI 0.88): (0) narrowed laterally; (1) widened laterally.

In the majority of Hemirhipini the metacoxal plate is narrowed laterally. In one group formed by Tetrigus parallelus and Pherhimius, Catelanus, Fusimorphus, Saltamartinus and Hemirhipus species, Alaomorphus candezei and Pseudocalais basilewskyi it is widened laterally.

54. Free margin of metacoxal plate (CI 0.25 RI 0.71): (0) straight; (1) with wide lobe; (2) with small lobe.

In the majority of Hemirhipini the free margin of metacoxal plate is straight. In Aphileus lucanoides, Tetrigus parallelus and one group formed by Pherhimius, Catelanus, Fusimorphus and Saltamartinus species it presents one wide lobe; in Eumoeus murray, Hemirhipus species, Conobajulus ugiensis, Cryptalaus prosectus and Alaomorphus candezei it presents one small lobe.

55. Scutellum (CI 0.25 RI 0.93): (0) horizontal or declivous; (1) angular.

In the majority of Hemirhipini the scutellum is horizontal or declivous. In Fusimorphus submetallicus, Hemirhipus species and one large group of Chalcolepidius species, viridipilis+, the scutellum is folded forming an angle.

56. Scutellum horizontal or declivous (CI 0.27 RI 0.58): (0) subhexagonal; (1) subpentagonal; (2) subquadrangular; (3) subrectangular; (4) vase-like; (5) triangular.

Among the Hemirhipini with scutellum horizontal or slightly declivous, the majority is sub-pentagonal in shape. In Eumoeus murray, Lacais species and Alaus lusciosus it is subhexagonal; in Anthracalaus westermanni and Eleuphemus and Propalaus species it is subquadrangular; in Aphileus lucanoides, Alaolacon cyanipennis, Phibisa pupieri, Alaus unicus, A. sericeus and A. zunianus, it is subrectangular; in Calais excavatus, a large group of Chalcolepidius species, supremus+, and C. rubripennis, it is vase-like; in Mocquerysia coerulipennis, it is triangular, representing an autapomorphy to this species.

57. Declivity of scutellum (CI 0.20 RI 0.60): (0) slight or moderate; (1) strong.

The declivity of not folded scutellum is variable and usually is slightly or moderately declivous. In two groups of Chalcolepidius species, extenuatuvittatus+ and mexicanus+, and C. rubripennis it is strongly declivous.

58. Horizontal area of scutellum angular (CI 1.00 RI 1.00): (0) elliptical; (1) subtrapezoidal; (2) rounded.

The shape of the horizontal area of angular scutellum is variable. In Hemirhipus species it is elliptical; in a large group of Chalcolepidius species, viridipilis+, subtrapezoidal, representing a synapomorphy shared by the species of this group; in Fusimorphus submetallicus, rounded, representing an autapomorphy to this species.

59. Posterior margin of scutellum (CI 0.17 RI 0.61): (0) strongly notched at middle; (1) slightly notched at middle; (2) prominent at middle; (3) rounded; (4) sinuous; (5) straight.

The shape of posterior margin of scutellum is variable among the Hemirhipini. It is strongly notched at middle in Lacais species, Abiphis nobilis, Coryleus pectinatus and Propalaus alicii; slightly notched at middle in Hemirhipus species, Calais excavatus, Phibisa pupieri, Lycoreus goudoti, Propalaus haroldi, Alaus tricolor, one large group of Chalcolepidius species, jansoni+, and some other species of this genus; prominent at middle in Aphileus lucanoides, Thoramus wakefieldi, Tetrigus parallelus, Pherhimius and Saltamartinus species, Aliteus reichei, Conobajulus ugiensis, Cryptalaus prosectus, one group formed by Neocalais macer and Austrocalais pogonodes, Pseudocalais basilewskyi, Chalcolepis species and some species of Alaus and some of Chalcolepidius; rounded in one group formed by Eumoeus murray, Alaolacon cyanipennis, Mocquerysia coerulipennis, Eleuphemus species, Fusimorphus submetallicus, Alaomorphus candezei, Alaus species (except A. tricolor, A. latipennis, A. patricius, A. calcaripilosus and A. thoracopunctatus), one group of Chalcolepidius species, supremus+; sinuous in Chalcolepidius desmaresti; straight in Anthracalaus westermanni, representing an autapomorphy to this species.

60. Wedge cell of hind wing (CI 1.00 RI 1.00): (0) present; (1) absent.

The wedge cell of hind wing is present only in Alaolacon cyanipennis, representing an autapomorphy to this species.

61. Number of sclerotizations at apex of hindwing (CI 0.50 RI 0.85): (0) one; (1) two, almost same size; (2) two, one longer; (3) three.

The hindwing of the majority of Hemirhipini presents two sclerotizations at apex,one longer than other. The hindwing of Aphileus lucanoides, Hemirhipus species, Lacais glauca and L. suturalis has one sclerotization; Anthracalaus westermanni, one group formed by Eumoeus murray, Alaolacon cyanipennis, Mocquerysia coerulipennis and Eleuphemus species, Thoramus wakefieldi, Pherhimius species and Aliteus reichei, two sclerotizations, both almost of the same size; Tetrigus parallelus, three sclerotizations, representing an autapomorphy to this species.

62. Lateral margins of elytra (CI 0.50 RI 0.00): (0) straight or slightly rounded; (1) strongly rounded.

The majority of the Hemirhipini has lateral margins of elytra straight or slightly rounded. In Aphileus lucanoides and Conobajulus ugiensis the lateral margins of elytra are strongly rounded.

63. Interstices (CI 0.17 RI 0.72): (0) strong or moderately convex; (1) alternate; (2) irregular; (3) flat.

The majority of the Hemirhipini presents elytra with interstices strong or moderately convex. The interstices are alternate in Catelanus trilineatus, Hemirhipus species, Propalalus haroldi and Chalcolepis species, Alaus veracruzanus, one group of Alaus species, cinnamomeus+, Chalcolepidius pruinosus, C. obscurus and C. silbermanni; variable in different areas of elytra considered as irregular, in Chalcolepidius forreri, representing an autapomorphy to this species; flat in Anthracalaus westermanni, Aphileus lucanoides, Mocquerysia coerulipennis, one group formed by Pherhimius species, Catelanus trilineatus, Fusimorphus submetallicus, Saltamartinus species, Conobajulus ugiensis, Austrocalais pogonoides, Alaomorphus candezei, one large group of Alaus species, sericeus+, Chalcolepidius lenzi, C. smaragdinus and one group of Chalcolepidius species, apacheanus+.

64. Third elytral interstice (CI 1.00 RI 1.00): (0) flat or convex in all length; (1) raised near base forming one tubercle; (2) raised near base forming one longitudinal ridge; (3) raised near base forming one dentiform tubercle; (4) expanded laterally forming one flattened tubercle.

The third interstice of elytrum is flat or convex in the majority of studied Hemirhipini. In Conobajulus ugiensis it is raised near base forming a tubercle, representing an autapomorphy to this species; in Austrocalais pogonodes it is raised near base forming a longitudinal ridge, representing an autapomorphy to this species; in Cryptalaus prosectus it is raised near base forming a dentiform tubercle, representing an autapomorphy to this species; in Alaomorphus candezei it is expanded laterally forming a flattened tubercle, representing an autapomorphy to this species.

65. Apex of elytrum (CI 0.33 RI 0.71): (0) rounded; (1) truncate.

The apex of elytrum of the majority of Hemirhipini is rounded. It is truncate in one group formed by Conobajulus ugiensis, Calais excavatus and Cryptalaus prosectus, Austrocalais pogonodes and one group of Alaus species, nobilis+.

66. Sutural spine of elytrum (CI 0.06 RI 0.40): (0) absent; (1) present.

The sutural spine of apex of elytrum is absent in the majority of Hemirhipini. It is present in Tetrigus parallelus, one group formed by Catelanus trilineatus and Fusimorphus submetallicus, Cryptalaus prosectus, one group formed by Neocalais macer and Austrocalais pogonodes, Coryleus pectinatus, Pseudocalais basilewskyi, Chalcolepis species, one group of Alaus species, nobilis+, A. thoracopunctatus and some Chalcolepidius species.

67. Lateroapical spine of elytrum (CI 0.33 RI 0.33): (0) absent; (1) present.

The lateroapical spine of apex of elytrum is absent in the majority of Hemirhipini. It is present in Calais excavatus, Austrocalais pogonodes and one group of Alaus species, latipennis+.

68. Tibial spurs (CI 0.33 RI 0.90): (0) present; (1) absent.

The tibial spurs are absent in the majority of Hemirhipini. It is present in Anthracalaus westermanni, Aphileus lucanoides, one group formed by Eumoeus murray, Alaolacon cyanipennis, Mocquerysia coerulipennis and Eleuphemus species, Thoramus wakefieldi, one group formed by Catelanus trilineatus, Fusimorphus submetallicus and Saltamartinus species and Hemirhipus species.

69. Basal setae of claws (CI 1.00 RI 1.00): (0) present; (1) absent.

All Hemirhipini present setae at base of claws.

70. Apex of last ventrite of female (CI 0.20 RI 0.89): (0) rounded; (1) truncate with fringe of spatulate setae.

The apex of last ventrite of female is rounded or truncate. When truncate, it is clothed with a dense fringe of spatulate setae. Apex truncate with fringe of spatulate setae is present in one group formed by Calais excavatus and Cryptalaus prosectus, one group formed by Neocalais macer, Austrocalais pogonodes, Lacais species, Abiphis nobilis, Phibisa pupieri, Coryleus pectinatus and Lycoreus goudoti, one group formed by Pseudocalais basilewskyi and Propalaus species, Alaus veracruzanus, A. unicus, A. tricolor, A. patricius, group calcaripilosus+, and Chalcolepidius species.

71. Anterior tibiae of male (CI 0.30 RI 0.89): (0) without ornamentations or with stout setae; (1) bearing spines; (2) bearing fringe of short cilia; (3) bearing fringe of long cilia.

The anterior tibiae of male do not present ornamentations in the majority of Hemirhipini. In Aphileus lucanoides, Propalaus alicii, Alaus species (except A. cinnamomeus) and one large group of Chalcolepidius species, supremus+, they present spines; in some (few) Chalcolepidius species they present a fringe of short cilia; in one large group of Chalcolepidius species, virginalis+, a fringe of long cilia.

72. Median tibiae of male (CI 0.50 RI 0.93): (0) without ornamentations or with stout setae; (1) bearing spines; (2) bearing fringe of long cilia; (3) bearing fringe of short cilia.

The median tibiae of male do not present ornamentations in the majority of Hemirhipini. In Aphileus lucanoides, Alaus calcaripilosus and one large group of Chalcolepidius species, supremus+, they present spines; in one large group of Chalcolepidius species, porcatus+, they present a fringe of long cilia; in Chalcolepidius tartarus, a fringe of short cilia, representing an autapomorphy to this species.

73. Fourth tarsomere (CI 1.00 RI 1.00): (0) simple; (1) lamellate.

In all studied Hemirhipini the fourth tarsomere are not lamellate.

74. Ventral region of last tarsomere of male (CI 0.75 RI 0.96): (0) with short setae; (1) with fringe of long cilia; (2) with fringe of short cilia; (3) with fringe of long cilia at distal half of tarsomeres.

In the majority of Hemirhipini the ventral region of last tarsomere of male is clothed with short setae. In one large group of Chalcolepidius species, virginalis+, they have a fringe of long cilia; in Chalcolepidius tartarus, a fringe of short cilia, representing an autapomorphy to this species; in Chalcolepidius fasciatus, a fringe of long cilia only at distal half of tarsomere, representing an autapomorphy to this species.

75. Shape of sternite 8 of male (CI 0.30 RI 0.76): triangular or trapezoidal; (1) subpentagonal; semielliptical; (3) band-like.

The shape of the sternite 8 of male is variable and in the majority of Hemirhipini it is band-like. In Tetrigus parallelus, Catelanus trilineatus, one group formed by Calais excavatus and Cryptalaus prosectus, one group formed by Neocalais macer, Lacais species, Alaomorphus candezei, Abiphis nobilis and Phibisa pupieri, Propalaus alicii, Alaus species (except A. patricius) and one group of Chalcolepidius species, desmaresti+, triangular or trapezoidal; in Lycoreus goudoti, Pseudocalais basilewskyi, Alaus patricius, Chalcolepidius tartarus and C. boucardi, semielliptical.

76. Distal margin of sternite 8 male (CI 0.18 RI 0.72): (0) notched; (1) strongly notched; (2) prominent at middle; (3) rounded.

The distal margin of sternite 8 of male is notched at middle in Anthracaclaus westermanni, Eumoeus murray, Pherhimius species, Catelanus trilineatus, Saltamartinus scriptus, Hemirhipus species, Propalaus alicii and some species of Chalcolepidius; strongly notched at middle in Eleuphemus funerarius, representing an autapomorphy to this species; prominent at middle in Tetrigus parallelus, Alaomorphus candezei, Chalcolepis species and several species of Chalcolepidius from groups supremus+ and desmaresti+; rounded in a group formed by Calais excavatus and Cryptalaus prosectus, one group formed by Neocalais macer, Lacais species, Abiphis nobilis, Phibisa pupieri, and Lycoreus goudoti, Pseudocalais basilewskyi and Alaus species.

77. Melanized area of sternite 8 of male (CI 0.20 RI 0.77): (0) entire; (1) broken; (2) absent.

The sternite 8 of male is translucent and presents one melanized band near margins.This area is entire when it runs near anterior, lateral and posterior margins without interruptions, present in Neocalais macer, Alaomorphus candezei, Abiphis nobilis, Alaus sericeus, A. plebejus, A. oculatus and one large group of Chalcolepidius species, desmaresti+. This area is broken or divided in parts in the majority of studied Hemirhipini; it is absent and whole sternite is translucent in Phibisa pupieri and Chalcolepidius mexicanus.

78. Melanized broken area of sternite 8 of male (CI 0.42 RI 0.70): (0) U-shaped; (1) divided in three parts; (2) broken median basally; (3) broken median distally; (4) U-inverted; (5) divided in two parts; (6) narrow transverse basal band.

When the melanized area is broken, it presents variable shapes. It is U-shaped when is absent at base, present in Tetrigus parallelus, Pherhimius fascicularis, one group formed by Calais excavatus and Crypatalus prosectus, Lacais species, one group formed by Pseudocalais basilewskyi, Propalaus alicii and Chalcolepis species, Alaus veracruzanus, A. unicus, A. tricolor and A. patricius; it is divided in three parts in Pherhimius dejeani, Alaus myops, A. lusciosus and some Chalcolepidius species of group ferratovittatus+; it is broken median basally in Catelanus trilineatus, Saltamartinus scriptus, Lycoreus goudoti and some Chalcolepidius species of group supremus+; it is broken median distally in Alaus melanops, representing an autapomorphy to this species; it is U-shaped inverted when it is absent near anterior margin, in Anthracalaus westermanni, Eumoeus murray, Hemirhipus species and Alaus zunianus; divided in two parts in Alaus calcaripilosus and A. thoracopunctatus; as one narrow transverse band near basal margin, in Eleuphemus funerarius.

79. Distal margin of tergite 9 of male (CI 0.27 RI 0.66): (0) slightly notched; (1) moderately notched; (2) strongly notched; (3) rounded.

The distal margin of tergite 9 of male is moderately notched in the majority of Hemirhipini. It is slightly notched in Thoramus wakefieldi, Catelanus trilineatus, Calais excavatus, Pseudocalais basilewskyi and Propalaus alicii; strongly notched in Anthracaclaus westermanni, Eumoeus murray, Eleuphemus funerarius and one group of Chalcolepidius species, desmaresti+; rounded in two groups of Chalcolepidius species, obscurus+ and aurulentus+.

80. Position of basal piece of aedeagus (CI 0.50 RI 0.97): (0) above parameres base; (1) dorsally, on basal fourth of parameres.

The basal piece of aedeagus is above the parameres base in the majority of studied Hemirhipini. It is dislocated dorsally on basal fourth of para-meres in Alaomorphus candezei and one large group of Chalcolepidius species, desmaresti+.

81. Parameres (CI 0.33 RI 0.66): (0) fused; (1) separated.

The parameres of aedeagus are fused in the majority of Hemirhipini. They are separated in basal groups, Anthracalaus westermanni, Aphileus lucanoides, Eumoeus murray, Mocquerysia coerulipennis, Thoramus wakefieldi and Tetrigus parallelus.

The presence of separated parameres was used by Casari-Chen (1994) to remove some genera from Hemirhipini. The presence of parameres fused was considered a synapomorphy to the tribe.

82. Parameres fused (CI 0.16 RI 0.73): (0) fused em large area; (1) fused in short area.

In the majority of Hemirhipini with parameres fused, the fusion occurs in a large area; fusion in a short area occurs in Eleuphemus funerarius, Catelanus trilineatus, Fusimorphus submetallicus, Saltamartinus scriptus, Hemirhipus species, Cryptalaus prosectus, Phibisa pupieri, Alaus unicus and one group of Alaus species, thoracopunctatus+, and A. oculatus.

83. Subapical region of parameres (CI 1.00 RI 1.00): (0) unciform without small teeth; (1) not unciform.

All studied Hemirhipini present subapical region of parameres not unciform, representing a synapomorphy shared by all tribe. This character had already been considered as a synapomorphy to tribe by Casari-Chen (1994).

84. Subapical region of parameres not unciform (CI 0.50 RI 0.83): (0) spearhead-like; (1) cleft laterally; (2) slightly securiform; (3) slightly securiform with dorsolateral tooth; (4) securiform; (5) securiform with tooth; (6) spatula like; (7) with uncus dorsal; (8) almost straight.

The subapical region of the parameres of the Hemirhipini aedeagus are of variable shapes, but never unciform. They are spearhead-like in a large group of Chalcolepidius species, supremus+, representing a synapomorphy shared by species of this group; cleft laterally in a very large group of Chalcolepidius species, desmaresti+; slightly securiform in Aphileus lucanoides, Cryptalaus prosectus, Lacais species, Alaus veracruzanus and A. plebejus; slightly securiform with dorsolateral tooth in Chalcolepis species, representing a synapomorphy shared by species of this genus; securiform in Eumoeus murray, and one group formed by Pherhimius species, Catelanus trilineatus, Fusimorphus submetallicus and Saltamartinus scriptus; securiform with tooth in Calais excavatus, Lycoreus goudoti, Pseudocalais basilewskyi, Propalaus alicii, two groups of Alaus species, cinnamomeus+ and thoracopunctatus+ and Alaus sericeus; spatula like in Thoramus wakefieldi and Hemirhipus species; with uncus dorsal in one group of Alaus species, nobilis+, representing a synapomorphy shared by species of this group; almost straight in Mocquerysia coerulipennis and Eleuphemus funerarius.

85. Lateral cleft of parameres (CI 0.66 RI 0.66): (0) moderately long; (1) short; (2) inverted.

The lateral cleft of parameres usually starts near apex and goes inclined upwards to median direction. The cleft is moderately long in a large group of Chalcolepidius species, desmaresti+, representing a synapomorphy shared by species of this group; short in Conobajulus ugiensis and Chalcolepidius boucardi. In Neocalais macer, Alaomorphus candezei and Pseudocalais basilewskyi the cleft is inverted: it is inclined downwards to median direction.

86. Apex of region delimited by lateral cleft of para-meres (CI 0.62 RI 0.57): (0) with truncate apex; (1) with rounded apex; (2) with excavated apex; (3) rounded in one angle; (4) with triangular apex; (5) preceeded by lobe.

The lateral cleft delimits a subapical area of parameres that presents apex of different shapes. It has truncate apex in a large group of Chalcolepidius species, viridipilis+, representing a synapomorphy shared by species of this group; rounded apex in a group of Chalcolepidius species, desmaresti+, and C. rubripennis and C. chalcantheus; excavated apex in Conobajulus ugiensis, Chalcolepidius virgatipennis and C. erythroloma; apex with one lateral angle rounded in Chalcolepidius virginalis and C. webbi; triangular apex in one group formed by Neocalais macer, Abiphis nobilis, Phibisa pupieri, Coryleus pectinatus, Lycoreus goudoti and Alaomorphus candezei, representing a synapomorphy shared by species of this group; cleft preceeded by lobe in Pseudocalais basilewskyi, representing an autapomorphy to this species.

87. Subapical region of area delimited by lateral cleft of parameres (CI 0.87 RI 0.85): (0) narrow with truncate apex; (1) narrow with rounded apex; (2) flat and concave; (3) spoon-like apex; (4) short with dorsal uncus; (5) wide; (6) triangular; (7) sharpened.

The area delimited by lateral cleft presents different shapes. It is narrow with truncate apex in Chalcolepidius inops and one group of Chalcolepidius species, apacheanus+; narrow with rounded apex in in a group of Chalcolepidius species desmaresti+, representing a synapomorphy to species of this group; narrow flat and concave in one group of Chalcolepidius species, virgatipennis+, representing a synapomorphy shared by this group of species; narrow with spoon-like apex in Chalcolepidius chalcantheus, representing an autapomorphy to this species; short with dorsal uncus in Chalcolepidius virginalis, representing an autapomorphy to this species; wide in one group of Chalcolepidius species, viridipilis+, representing a synapomorphy shared by species of this group; triangular in Neocalais macer, Alaomorphus candezei and Pseudocalais basilewskyi; sharpened in Conobajulus ugiensis, representing an autapomorphy to this species.

88. Basal region of median lobe (CI 0.33 RI 0.66): (0) straight or slightly rounded; (1) constricted; (2) with tooth or lobe.

The aedeagus of the majority of the Hemirhipini presents basal region of median lobe straight or slightly rounded. Basal region of median lobe constricted is present in one group of Alaus species, cinnamomeus+, and A. tricolor; with tooth or lobe in Pherhimius and Hemirhipus species and Pseudocalais basilewskyi.

89. Basal region of median lobe without constriction (CI 1.00 RI 1.00): (0) without ornamentations; (1) with lobe; (2) with tooth and lobe.

When median lobe of aedeagus is not constricted, usually it does not present ornamentation. It presents one lobe each side in Hemirhipus species (except H. elegantissimus), representing a synapomorphy shared by species of this genus, and tooth and lobe in Hemirhipus elegantissimus, representing an autapomorphy to this species.

90. Median lobe (CI 0.46 RI 0.68): (0) gradually narrowed apicad; (1) constricted near middle; (2) narrowed near middle; (3) narrowed near base; (4) almost straight; (5) widened in distal half; (6) widened near base.

The median lobe of Hemirhipini presents different shapes. It is gradually narrowed apicad in Anthracalaus westermanni, Aphileus lucanoides, Eumoeus murray, Mocquerysia coerulipennis, Eleuphemus funerarius, Thoramus wakefieldi, Tetrigus parallelus, Pherhimius species, Saltamartinus scriptus, one group formed by Calais excavatus and Cryptalaus prosectus, one group formed by Neocalais macer, Lacais species, Alaomorphus candezei, Abiphis nobilis and Phibisa pupieri, Propalaus alicii, Alaus species (except A. calcaripilosus, A. myops and A. oculatus), one large group of Chalcolepidius species, desmaresti+, and some other species of this genus; constricted near middle in Lycoreus goudoti, one group of Chalcolepidius species, mexicanus+, and C. angustus; narrowed near middle in Conobajulus ugiensis and Chalcolepis species; narrowed near base in Catelanus trilineatus and Pseudocalais basilewskyi; almost straight in Fusimorphus submetallicus, representing an autapomorphy to this species; widened at distal half in Alaus calcaripilosus, representing an autapomorphy to this species; widened near base in Hemirhipus species, Alaus myops and A. oculatus.

91. Median lobe (CI 0.66 RI 0.97): (0) without lateral teeth; (1) with lateral teeth; (2) with dorsolateral small teeth.

In the majority of Hemirhipini, the median lobe of aedeagus is not toothed laterally. It presents lateral teeth in Pseudocalais basilewskyi, Propalaus alicii and one very large group of Chalcolepidius species, desmaresti+; small dorsolateral teeth in Chalcolepidius boucardi, representing an autapomorphy to this species.

92. Number of teeth of median lobe (CI 0.33 RI 0.33): (0) under 20; (1) above 30.

The Chalcolepidius species of group desmaresti+ present median lobe toothed laterally with about 20 teeth each side; median lobe with more than 30 teeth each side is found in Pseudocalais basilewskyi, Propalaus alicii, Chalcolepidius jansoni and C. viridipilis.

93. Width of median lobe base/parameres (CI 0.16 RI 0.59): (0) 0.35-0.44; (1) 0.45-0.50; (2) 0.51-0.60; (3) 0.61-0.70; (4) 0.71-0.80.

The relation between the width of the median lobe base and the width of the parameres, at same point, varies from 0.35 to 0.44 in Thoramus wakefieldi, Pherhimius fascicularis, Lacais species, Alaomorphus candezei, Propalaus alicii, Alaus lusciosus and a very large group of Chalcolepidius species, desmaresti+; from 0.45 to 0.50 in Eleuphemus funerarius, one group formed by Tetrigus parallelus, Pherhimius dejeani, Catelanus trilineatus, Saltamartinus and Hemirhipus species, Neocalais macer, Phibisa pupieri, Alaus sericeus, two groups of Alaus species, calcaripilosus+ and melanops+, one group of Chalcolepidius species, ferratuvittatus+, C. exulatus and C. serricornis; from 0.51 to 0.60 in Fusimorphus submetallicus, one group of Hemirhipus species, apicalis+, Calais excavatus, one group formed by Pseudocalais basilewskyi, Chalcolepis species and Alaus species, and a group of Chalcolepidius species, mexicanus+; from 0.61 to 0.70 in Aphileus lucanoides, Eumoeus murray, Conobajulus ugiensis, Cryptalaus prosectus, one group formed by Abiphis nobilis, Phibisa pupieri, Coryleus pectinatus and Lycoreus goudoti, Alaus unicus, A. patricius and Chalcolepidius angustatus; from 0.71 to 0.80 in Anthracalaus westermanni and Alaus tricolor.

94. Sternite 8 of female (CI 0.80 RI 0.93): (0) triangular; (1) subquadrangular or transverse; (2) pentagonal; (3) rounded; (4) trapezoidal.

The sternite 8 of female in the majority of Hemirhipini is subquadrangular or transverse. The sternite 8 triangular represents one synapomorphy shared by majority of Hemirhipini, excluding only the more basal clades representing the genera Anthracalaus, Aphileus and one group representing Eumoeus, Alaolacon, Mocquerysia and Eleuphemus. Sternite subquadrangular represents one synapomorphy for a large group of Hemirhipini genera: Conobajulus, Calais, Cryptalaus, Neocalais, Austrocalais, Lacais, Alaomorphus, Abiphis, Phibisa, Coryleus, Lycoreus, Pseudocalais, Propalaus, Chalcolepis, Alaus and Chalcolepidius.

Sternite pentagonal in Aphileus lucanoides and Eleuphemus fasciatus; rounded in a group formed by Alaolacon cyanipennis and Mocquerysia coerulipennis and Thoramus wakefieldi.

95. Sternite pentagonal in Aphileus lucanoides and Eleuphemus fasciatus; rounded in a group formed by Alaolacon cyanipennis and Mocquerysia coerulipennis and Thoramus wakefieldi. Sternite 8 of female (CI 0.20 RI 0.0): (0) wider than long; (1) longer than wide or as long as wide.

In the majority of Hemirhipin the sternite 8 of female are wider then long. In Mocquerysia coerulipennis, Eleuphemus fasciatus and Thoramus wakefieldi it is longer than wide or as long as wide.

96. Median region of distal margin of sternite 8 of female (CI 0.50 RI 0.88): (0) notched; (1) rounded; (2) prominent at middle.

In the majority of studied Hemirhipini the distal margin of sternite 8 of female is notched. It is rounded in one group formed by Alaolacon cyanipennis and Mocquerysia coerulipennis and Thoramus wakefieldi; prominent at middle in Aphileus lucanoides, Eleuphemus fasciatus and one group formed by Tetrigus parallelus, Pherhimius species, Catelanus trilineatus, Fusimorphus submetallicus, Saltamartinus and Hemirhipus species and Aliteus reichei.

97. Distal margin of sternite 8 of female notched (CI 0.22 RI 0.55): (0) widely and strongly notched; (1) widely and moderately notched; (2) widely and slightly notched; (3) narrow and strongly notched; (4) narrow and moderately notched; (5) narrow and slightly notched.

When distal margin of sternite 8 of female is notched, the identation presents different sizes. It is is wide and deep in a group of Alaus species, calcaripilosus+, and A. unicus, one group of Chalcolepidius species, chalcantheus+, C. forreri and C. mocquerysi; wide and moderately deep in Lacais species, Alaus patricius, one large group of Chalcolepidius species, desmaresti+, and C. oxydatus; wide and slightly deep in Cryptalaus prosectus, one group formed by Neocalais macer and Austrocalais pogonodes, one group formed by Pseudocalais basilewskyi, Propalaus, Chalcolepis species (except C. luczotii) and Alaus species, one group of Chalcolepidius species, eschscholtzi+, C. inops, C. cyaneus and C. fryi; narrow and strongly deep in Calais excavatus, Phibisa pupieri, Lycoreus goudoti and one group of Chalcolepidus species, mexicanus+, C. extenuatuvittatus; narrow and moderately deep in Abiphis nobilis, Alaus latipennis and Chalcolepidius dugesi; narrow and slightly deep in Coryleus pectinatus, Chalcolepis luczotii and Alaus zunianus.

98. Relation width X spiculum gastrale of sternite 8 of female (CI 0.18 RI 0.62): (0) 0.25-0.50; (1) 0.51-0.75; (2) 0.76-1.0; (3) 1.01-1.25; (4) 1.26-1.50.

The relation between the width of the sternite 8 of female and the length of the spiculum gastrale varies from 0.25 to 0.50 in Tetrigus parallelus, Fusimorphus submetallicus, Hemirhipus species and Alaus melanops; from 0.51 to 0.75 in Mocquerysia coerulipennis, Eleuphemus fasciatus, Thoramus wakefieldi, Pherhimius species, Catelanus trilineatus, Aliteus reichei, a groups formed by Calais excavatus and Cryptalaus prosectus, Chalcolepis esplendidus, Alaus latipennis and a group of Alaus species, thoracopunctatus+; from 0.76 to 1.0 in Alaolacon cyanipennis, Austrocalais pogonodes, Lacais nietoi, one group formed by Phibisa pupieri, Coryleus pectinatus and Lycoreus goudoti, Propalaus haroldi, Chalcolepis species (except C. splendidus), Alaus unicus, A. tricolor, A. patricius and A. plebejus, two groups of Chalcolepidius species, supremus+ (except albisetosus+) and desmaresti+ (except chalcantheus+ and inops+) from 1.01 to 1.25 in Aphileus lucanoides, Neocalais macer, a group of Lacais species, glauca+, Abiphis nobilis, three groups of Chalcolepidius species, albisetosus+, chalcantheus+ and inops+, C. gossipiatus, C. serricornis, C. spinipennis, C. rubripennis, C. zonatus, C. mocquerysii and C. rugatus; from 1.26 to 1.50 in Alaus calcaripilosus and one group of Chalcolepidius species, boucardi+.

99. Relation sternite 8 X spiculum gastrale length (CI 0.15 RI 0.51): (0) 0.20-0.40; (1) 0.41-0.6; (2) 0.61-0.80; (3) 0.81-1.0; (4) 1.01-1.20; (5) 1.21-1.40; (6) 1.41-160; (7) 1.61-180; (8) 1.81-2.0; (9) above 2.1.

The relation between the length of the sternite 8 of female and the length of the spiculum gastrale varies from 0.20 to 0.40 in one group formed by Tetrigus parallelus, Catelanus trilineatus, Fusimorphus submetallicus, Saltamartinus and Hemirhipus species and Alaus melanops; from 0.41 to 0.6 in one group formed by Alaolacon cyanipennis and Mocquerysia coerulipennis, Thoramus wakefieldi, Pherhimius species, Aliteus reichei, one group formed by Conobajulus ugiensis, Calais excavatus and Cryptalaus prosectus, one group formed by Neocalais macer, Austrocalais pogonodes, Lacais species, Alaomorphus candezei, Abiphis nobilis, Phibisa pupieri and Lycoreus goudoti, one group formed by Pseudocalais basilewskyi, Propalaus, Chalcolepis and Alaus species and several Chalcolepidius species, especially from groups mexicanus+ and desmaresti+; from 0.61 to 0.80 in Coryleus pectinatus and several Chalcolepidius species, especially from groups mexicanus+ and desmaresti+; from 0.81 to 1.0 in Aphileus lucanoides, Eleuphemus fasciatus, Chalcolepidius virgatipennis and C. eschscholtzi.

100. Ovipositor (CI 0.50 RI 0.42): (0) "normal"; (1) short and wide; (2) cuneiform; (3) long; (4) short and wide with dilated apex.

The ovipositor is considered "normal" when it is long, with about four times the coxites length, almost the same width as vagina and the coxites have rounded apex. This kind of ovipositor is present in the majority of studied Hemirhipini. It is short and wide in n Pherhimius fascicularis, Alaus calcaripilosus, Chalcolepidius chalcantheus and C. smaragdinus, it is about two time the coxites length, very wider than ovipositor and coxites with rounded apex; in one groups of Alaus species, thoracopunctatus+, it is long with coxites cuneiform, representing a synapomorphy shared by species of this group; in Aphileus lucanoides it is short and wide with dilated apex, representing an autapomorphy to this species.

101. Ovipositor (CI 1.00 RI 1.00): (0) without styli; (1) with styli.

All studied Hemirhipini present ovipositor without styli.

102. Sclerotized pieces of openings of colleterial glands (CI 0.25 RI 0.76): (0) present; (1) absent.

The majority of studied Hemirhipini presents two sclerotized pieces at openings of colleterial glands. They are absent in Mocquerysia coerulipennis, one group formed by Thoramus wakefieldi, Tetrigus parallelus, Pherhimius species, Catelanus trilineatus, Fusimorphus submetallicus, Saltamartinus and Hemirhipus species.

103. Shape of sclerotized pieces of openings of colleterial glands (CI 0.57 RI 0.92): (0) elliptical and opened; (1) elliptical and closed; (2) elongate; (3) funil-like; (4) triangular.

The shapes of the sclerotized pieces are variable and in the majority of Hemirhipini are elliptical and opened laterally. They are elliptical and closed in Pherhimius dejeani and one large group of Chalcolepidius species desmaresti+; they are elongate in Eleuphemus fasciatus and Alaus tricolor; funil-like in Anthracalaus westermanni, representing an autapomorphy to this species; triangular in Aphileus lucanoides, representing an autapomorphy to this species.

104. Sclerotized piece of openings of colleterial glands (CI 0.37 RI 0.56): (0) without teeth; (1) with one row of teeth; (2) with some rows of teeth at base; (3) with several rows of teeth in 1/3; (4) with many rows of teeth in 1/2 of diameter; (5) greater-like; (6) with small teeth.

The sclerotized pieces at openings of colleterial glands do not have teeth in Anthracalaus westermanni, Aphileus lucanoides, Lacais species (except L. glauca), Chalcolepis species (except C. austerus), Alaus thoracopunctatus and one group of Chalcolepidius species, supremus+. The majority of species has one row of teeth that represents a synapomorphy to a large group of Hemirhipini that excludes only Anthracalaus and Aphileus. Sclerotized pieces with some rows of teeth at base is present in Chalcolepidius porcatus, C. silbermanni and C. rostainei; with several rows of teeth disposed in 1/3 of the area of the pieces in Chalcolepidius chalcantheus and C. boucardi; with many rows of teeth disposed in ½ of the diameter of the pieces in Chalcolepidius viridipilis, C. tartarus, C. smaragdinus and C. inops; with many small teeth, greater-like, in Chalcolepidius desmaresti, representing an autapomorphy to this species; with several small teeth in Pherhimius dejeani, representing an autapomorphy to this species.

105. Bursa copulatrix (CI 0.22 RI 0.88): (0) elongate; (1) elongate, very narrow; (2) rounded.

The bursa copulatrix is elongate and moderately wide in Anthracalaus westermanni, Aphileus lucanoides, Calais excavatus, Lacais species, one group formed by Coryleus pectinatus and Lycoreus goudoti, Pseudocalais basilewskyi, Chalcolepis and Alaus species, one group of Chalcolepidius species, supremus+ (except rubripennis+) several species not observed; C. desmaresti and C. jansoni); elongate and very narrow in Eleuphemus fasciatus, one group formed by Thoramus wakefield, Tetrigus parallelus, Pherhimius species, Catelanus trilineatus, Fusimorphus submetallicus, probably Saltamartinus and Hemirhipus species, Aliteus reichei and Propalaus haroldi; rounded in Mocquerysia coerulipennis, Cryptalaus prosectus, one group formed by Neocalais macer and Austrocalais pogonodes, one group formed by Abiphis nobilis and Phibisa pupieri and one group of Chalcolepidius species, rubripennis+.

106. Spiny areas of bursa copulatrix (CI 0.20 RI 0.42): (0) present; (1) absent.

The bursa copulatrix of the majority of Hemirhipini presents some spiny areas. In Aphileus lucanoides, Mocquerysia coerulipennis, Eleuphemus fasciatus, Aliteus reichei, group formed by Calais excavatus and Cryptalaus prosectus, Alaus latipennis and A. oculatus bursa copulatrix does not have spines.

107. Number of spiny areas of bursa copulatrix (CI 0.31 RI 0.71): (0) one longitudinal; (1) one longitudinal and two rounded lateral; (2) totally spiny ventrally or dorsally; (3) small dorsal area; (4) totally spiny; (5) two spiny areas; (6) one longitudinal dorsal covering all basal area.

Bursa copulatrix with one longitudinal and elongate spiny area is present in Chalcolepis austerus, C. similis, Alaus species (except A. veracruzanus, A. cinnamomeus, A. sericeus, A. nobilis, A. latipennis, A. patricius and A. oculatus), Chalcolepidius extenuatuvittatus, C. gossipiatus, C. rubripennis and C. inops; one longitudinal and two lateral rounded spiny areas in Lacais glauca, a group of Chalcolepis species, luczotii+, two groups of Chalcolepidius species, desmaresti+ and virginalis+, C. mexicanus and C. serricornis; totally spiny, dorsally and ventrally in Coryleus pectinatus, Pseudocalais basilewskyi, Alaus patricius, one group of Chalcolepidus species, attenuatus+ and C. spinipennis; spiny in one dorsal small area in Lacais nietoi, one group of Chalcolepidius species, viridipilis+, and C. dugesi; totally spiny in Anthracalaus westermanni, Thoramus wakefieldi, Pherhimius species, one group formed by Catelanus trilineatus and Fusimorphus submetallicus, Hemirhipus species, Austrocalais pogonodes, Lacais suturalis, one group formed by Abiphis nobilis and Phibisa pupieri and Lycoreus goudoti; two spiny areas in Tetrigus parallelus, representing an autapomorphy to this species; one longitudinal dorsal spiny area covering all basal area in Neocalais macer and Propalaus haroldi.

RESULTS AND DISCUSSION

The cladistic analysis resulted in two equally parsimonious trees. Both trees have CI 30, RI 74 and length = 924. The only difference between the two topologies is the position of Chalcolepidius corpulentus: in one tree it is the sister-group of limbatus+ and in other, it forms a polytomy with ((C. approximatus) (C. bomplandi) (C. rostainei)) (limbatus+)). The consensus tree is represented by Figs. 1A, 1B.

Based on the cladistic analysis, the position of Alaolacon, Aliteus, Anthracalaus, Eumoeus, Mocquerysia and Tetrigus, removed from the tribe and not included in any suprageneric group, was corroborated inside the tribe Hemirhipini. Besides, it is demonstrated that Aphileus and Thoramus also belong to this tribe.

Casari (1996) presented a cladistic analysis to Alaus species, but in (2003) she included seven species in this genus. The present analysis revealed that the genus Alaus does not make a monophyletic group, and two species, Alaus alicii and A. haroldi, are more related to Pseudocalais basilewskyi than to other Alaus species. Propalaus gen. nov. is established to include these two species.

Based on the present analysis, the tribe Hemirhipini is characterized by synapomorhies: 20(2) nasal absent; 29(1) luminescent vesicles absent; 46(1) apex of prosternal spine rounded; 83(1) subapical region of parameres of aedeagus not unciform. The characters 46(1) and 83(1) were used by Casari-Chen (1994). It is formed by 30 genera: Abiphis, Alaolacon, Alaomorphus, Alaus, Aliteus, Anthracalaus, Aphileus, Austrocalais, Calais, Catelanus, Chalcolepidius, Chalcolepis, Conobajulus, Coryleus, Eleuphemus, Eumoeus, Fusimorphus, Hemirhipus, Lacais, Lycoreus, Mocquerysia, Neocalais, Paracalais, Pherhimius, Phibisa, Propalaus gen. nov., Pseudocalais, Saltamartinus, Tetrigus, Thoramus.

The Propalaus gen. nov. is the sister-group of Pseudocalais basilewskyi and it is defined by homoplasies: 23(1) antennae serrate or pectinate from 4^th antennomere; 44(1) prosternal channel present; 49(4) borders of mesosternal cavity horizontal and strongly declivous frontally; 56(2) scutellum subquadrangular. Besides, pubescence white ventrally, median anterior margin of pronotum raised forming two teeth and interstices alternate.

Propalaus gen. nov.

(Figs. 32, 33)

Type-species: Chalcolepidius haroldi Candèze, 1878.

Pubescence scale-like, dorsally black or whitish with brown small spots and narrow stripes and white ventrally. Frons not carinate. Antennae pectinate or strongly serrate; 3rd antennomere transverse, prominent laterally. Pronotum with lateral margins straight; strongly convex longitudinal medially; anterior margin with two small tubercles or two small teeth near middle; hind angles divergent and not carinate. Prosternal spine compressed laterally with cuneiform apex. Notosternal sutures straight; prosternal channel present. Borders of mesosternal cavity horizontal and strongly declivous frontally. Scutellum strongly declivous with posterior margin notched. Anterior tibiae with small spines; tibial spurs absent. Free margin of metacoxal plate straight. Interstices alternate; apices of elytra conjointly rounded.

This genus includes two species: P. alicii (Pjatakowa, 1941) and P. haroldi (Candèze, 1878).

- Integument black, median region of pronotum and interstices wine-red; pubescence black dorsally and white ventrally; anterior margin of pronotum forming two small tubercles near middle; mesosternal cavity U-shaped; meso-metasternal suture weak. Bolivia. (Fig. 32) ........................ P. alicii

- Integument black or reddish-brown; pubescence grayish- or yellowish-white; pronotum with two small rounded brown patches; higher interstices with sparser and brownish pubescence; anterior margin of pronotum forming two tubercles near middle; mesosternal cavity V-shaped; meso-metasternal suture well defined. Ecuador. (Fig. 33)........................ P. haroldi

A key to the genera of Hemirhipini

1. Antennae of male with 12 antennomeres and female with 11.................... 2 1'. Antennae with 11 antennomeres in both sexes..................................... 13 2(1). Antennae biflabellate in male and pectinate in female.............................. 3 2'. Antennae flabellate in both sexes or only in male.................................... 5 3(2). Integument black; pubescence scale-like wide, ashen with black spots on pronotum and elytra; subapical tooth of mandible absent; frons slightly carinate; pronotum moderately punctuate; hypomeral region strongly grooved near prosternal sutures; lateral margins of pronotum straight, gradually narrowed anteriad from hind angles apex; hind angles of pronotum backwards or slightly divergent and not carinate (Fig. 16). Senegal, Guinea, Ivory Coast, Sierra Leone, Benin, Central African Republic, Congo, Gabon, Tanzania, Democratic Republic of Congo, Angola, Mozambique..................... .................................................................... Eleuphemus Hyslop, 1921 3'. Integument dark- or reddish-brown; pubescence simple and thin; frons not carinate or slightly carinate; subapical tooth of mandible present; pronotum densely and coarsely punctuated; hypomera not grooved near prosternal sutures; lateral margins of pronotum straight with hind angles strongly divergent or slightly sinuous; hind angles of pronotum divergent and carinate........................................................................................ 4 4(3'). Integument dark-brown with hind angles of pronotum, antennae and legs reddish-brown; frons not carinate or slightly carinate; prothorax strongly narrower than elytra; pronotum grooved longitudinal medially; lateral margins of pronotum straight; hind angles strongly divergent; prosternal channnel present; prosternum moderately convex from sutures; median basal tubercle triangular elongate or indistinct; free margin of metacoxal plate straight; borders of mesosternal cavity horizontal and strongly declivous frontally; scutellum triangular; interstices flat (Fig. 23). Madagascar......................... ................................................................ Mocquerysia Fleutiaux, 1899 4'. Integument reddish-brown; frons not carinate; prothorax as wide or slightly narrower than elytra; pronotum grooved longitudinal medially at base; lateral margins of pronotum slightly sinuous; hind angles of pronotum backwards or slightly divergent; prosternal channel absent; prosternum strongly convex from sutures; median basal tubercle transverse; free margin of metacoxal plate with small lobe; borders of mesosternal cavity excavate at middle; scutellum subpentagonal; interstices convex (Fig. 17). India, Indonesia........ ..................................................................... Eumoeus Candèze, 1874 5(2'). Antennae flabellate in both sexes; pubescence simple or scale-like and velvet-like................................................................................................ 6 5'. Antennae flabellate in male and pectinate in female; pubescence scale-like or scale-like and velvet-like.................................................................. 10 6(5). Body fusiform; pubescence simple, fine; integument bright; unicolor or from yellow to ferrugineous with back spots or bands; sutural spine of elytra present......................................................................................... 7 6'. Body elliptical; pubescence scale-like and velvet-like or simple and thick, colorful and covering integument color; sutural spine of elytra absent........ 8 7(6). Unicolor; integument from dark-brown to black, with metallic bright; prothorax narrower than elytra; apex of hind angles of pronotum constricted; hind angles of pronotum carinate; borders of mesosternal cavity horizontal; scutellum folded (Fig. 18). French Guiana, Brazil....................................... ................................................................ Fusimorphus Fleutiaux, 1942 7'. Bicolor; integument bright with glabrous appearance, yellowish or ferrugineous with longitudinal black bands on pronotum and elytra; prothorax as wide as elytra; apex of hind angles straight; hind angles of pronotum not carinate; borders of mesosternal cavity declivous; scutellum declivous (Fig. 11). Brazil...................................................... Catelanus Fleutiaux, 1942 8(6'). Integument black and/or ferrugineous/red and black; general pubescence velvet-like and scale-like; pubescence forming longitudinal elliptical bands on pronotum, not reaching hind angles; frontal carina incomplete or absent; basal region of frons bearing median tubercle; notosternal sutures straight and slightly curved near apex; free margin of metacoxal plate with small lobe; scutellum folded or angular; interstices alternate (Fig. 21). Mexico, Guatemala, Colombia, Venezuela, Peru, French Guiana, Brazil (from Amazonas to Santa Catarina), Argentina, Uruguay.............. Hemirhipus Latreille, 1825 8'. Integument different from above; general pubescence simple, thick, long and dense; frons carinate; basal region of frons smooth; notosternal sutures straight; free margin of metasternal plate with wide lobe; scutellum declivous; interstices flat................................................................... 9 9(8'). Integument reddish- and yellowish-brown; elytra with sinuous transverse bands giving a marbled appearance; hind angles of pronotum with bifid carina; borders of mesosternal cavity horizontal in whole length; tibial spurs absent (Fig. 25). Southeast of United States of America, Mexico, Dominican Republic, Belize, Panama, French Guiana, Brazil (from Amazonas to Santa Catarina), Paraguay, Argentina, Uruguay............ Pherhimius Fleutiaux, 1942 9'. Integument dark-brown, reddish-brown and yellow; elytra with longitudinal elliptical basal and transverse apical bands; borders of mesosternal cavity excavate at middle; hind angles of pronotum unicarinate; tibial spurs present (Fig. 28). Mexico, Nicaragua, Suriname, Brazil........................................... .................................................................. Saltamartinus Casari, 1996 10(5'). Integument black; scale-like pubescence thick, white with black spots at base of pronotum, sides and apices of elytra; pubescence of sternites with white patches; lateral margins of pronotum rounded; prosternum widened at median region; notosternal sutures semicircular; prosternal channel absent; basal region of prosternal spine bearing longitudinal elongate groove; metasternum with wide groove parallel mesocoxal cavity; free margin of metacoxal plate with small lobe; elytra striate only basally; third interstice expanded laterally forming a flattened tubercle (Fig. 4). Kenya................... ................................................................ Alaomorphus Hauser, 1900 10'. Integument and pubescence different from above, never with scale-like pubescence thick; pubescence of sternites unicolor; lateral margins of pronotum sinuous; prosternum gradually widened anteriad; notosternal sutures straight; prosternal channel present or absent; prosternal spine without groove; metasternum without groove parallel metacoxal cavity; free margin of metacoxal plate straight; elytra totally striate; 3rd interstice different from above..................................................................... 11 11(10'). Integument black, reddish-brown or ferrugineous; pubescence scale-like, black and white; thorax long in relation to abdomen; convexity of pronotum gradual; median basal tubercle of pronotum transverse; prosternal channel absent; borders of mesosternal cavity declivous; scutellum subhexagonal (Figs. 19, 20). Mexico, Guatemala, Costa Rica, Nicaragua, Belize, Panama, Colombia, Suriname, French Guiana, north of Brazil, Peru............................ ........................................................................ Lacais Fleutiaux, 1942 11'. Integument and pubescence different form above; convexity of pronotum forming longitudinal median ridge, with one longitudinal groove each side; median basal tubercle of pronotum elongate; prosternal channel present; borders of mesosternal cavity horizontal; scutellum of other shape.......... 12 12(11'). Bicolor, integument from orange to ferrugineous with black spots or bands; pronotal integument forming median elliptical spot, covered by velvet-like pubescence; general pubescence velvet-like and scale-like; pubescence scale-like ferrugineous and velvet-like black, accompaining integument color; elytra striped with or without lateral bands; frons not carinate, not forming fold; meso-metasternal suture present; scutellum subpentagonal (Fig. 2). Madagascar....................................................... Abiphis Fleutiaux, 1926 12'. Unicolor, integument black or ferrugineous, without spots or bands; pronotum only with scale-like pubescence, fine, grayish or yellowish; frons not carinate and forming a rounded fold; meso- metasternal suture absent or obsolete; scutellum subrectangular (Fig. 26). Comoro Islands, Mozambique................. ........................................................................ Phibisa Fleutiaux, 1942 13(1'). Antennae flabellate in both sexes...................................................... 14 13'. Antennae serrate, strongly serrate or pectinate................................... 16 14(13). Unicolor, integument from red- to dark-brown; pubescence simple and dense; body narrow, almost parallel; frons incompletely carinate; 3rd antennomere simple, tranverse; lateral margins of pronotum slightly rounded; pronotum strongly convex from sutures; median basal tubercle of pronotum transverse; notosternal sutures slightly sinuous; elytra gradually narrowed at apex with sutural spine (Fig. 29). Japan, China, Korea, Philippines, India, Sri Lanka, Turkey, Lebanon, Sumatra, New Guinea, Australia..................................... ...................................................................... Tetrigus Candèze, 1857 14'. Unicolor or bicolor, integument of varied colors; pubescence scale-like or scale-like and velvet-like; body elliptical; frons not carinate; 3rd antennomere with spiniform appendix; lateral margins of pronotum sinuous; pronotum forming a longitudinal median ridge; median basal tubercle of pronotum elongate; elytra with apex truncate and rounded; sutural spine of elytra absent or very small............................................................... 15 15(14'). Integument totally black or with reddish elytra; pubescence scale-like, fine and greysh; nasal sloped; hind angles of pronotum backwards or slightly divergent; free margin of metacoxal plate straight; elytra with apex widely rounded with small sutural spine (Fig. 14). Comoro Islands.......................... ..................................................................... Coryleus Fleutiaux, 1942 15'. Integument black; pubescence scale-like and velvet-like, black and white; pronotum with velvet-like pubescence making a large discal spot divided or not at middle; nasal very short; hind angles of pronotum divergent; free margin of metacoxal plate with small lobe; elytra with apex widely rounded without sutural spine (Fig. 22). Madagascar........................................... .................................................................... Lycoreus Candèze, 1857 16(13'). Pubescence simple; unicolor; subapical tooth of mandible present........... 17 16'. Pubescence scale-like or scale-like and velvet-like; unicolor or bicolor; subapical tooth of mandible absent.................................................... 20 17(16). Mandibles very long, prominent; 3rd antennomere elongate; lateral margins of pronotum rounded with hind angles strongly divergent; interstices flat; lateral margin of elytra strongly rounded (Fig. 8). Australia................................... ...................................................................... Aphileus Candèze, 1857 17'. Mandibles not prominent; 3rd antennomere of other shape; lateral margins of pronotum straight or slightly rounded with hind angles slightly divergent or backwards; interstices convex or flat; lateral margins of elytra straight or slightly rounded.............................................................................. 18 18(17'). Integument brownish or dark-brown; frons carinate; nasal high; borders of mesosternal cavity horizontal and slightly declivous frontally; scutellum subpentagonal; tarsi long, usually longer than tibiae (Fig. 30). New Zealand............................................................. Thoramus Sharp, 1877 18'. Integument different from above; frons not carinate or incompletely carinate; nasal absent; scutellum of other shape; tarsi of moderate length, shorter than tibiae................................................................................... 19 19(18'). Elliptical; integument black; elytra with blue or green metallic bright; frons not carinate; prosternum strongly convex, flat longitudinally and grooved laterally; prosternal channel absent; pronotum grooved longitudinal medially near base; scutellum subrectangular; posterior margin of scutellum rounded; wedge cell of hind wing present; elytra slightly narrowed to apex; interstices convex (Fig. 3). Thailand, Malaysia...................... Alaolacon Candèze, 1865 19'. Fusiform; integument black and bright; frons incompletely carinate; prosternum strongly convex from sutures; prosternal channel present; pronotum moderately convex, not grooved longitudinalmedially; scutellum subquadrangular; posterior margin of scutellum straight; elytra gradually narrowed to apex; interstices flat (Fig. 7). China, Laos, Vietnam, Java, New Guinea, Australia......................................... Anthracalaus Fairmaire, 1888 20(16'). Pubescence velvet-like and scale-like; prosternal channel absent............ 21 20'. Pubescence only scale-like; prosternal channel present or absent............ 22 21(20). Pubescence scale-like of varied coloration, black, white, yellow and several tonalities of brown; velvet-like or scale-like pubescence forming two elliptical or rounded spots on pronotum, of varied sizes; prosternal channel absent; median basal tubercle transverse (Fig. 5). Canada, United States of America, Mexico, Bahamas, Cuba, Jamaica, Dominican Republic, Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, French Guiana, Brazil, Peru............................................. Alaus Eschscholtz, 1829 21'. Body elongate with sides almost parallel; pubescence scale-like of several tonalities of brown; pronotum with median elliptical spot dark-brown; frons incompletely carinate; nasal high; lateral margins of pronotum straight; pronotum stronger convex medioanteriorly; medioanterior margin of pronotum raised forming two teeth; prosternum widened frontally; metacoxal plate widened laterally; apex of elytra truncate with lateral angles rounded and sutural spine well developed (Fig. 27). Cameroon, Central African Republic, Democratic Republic of Congo, Tanzania, Gabon, Angola............................ .................................................................... Pseudocalais Girard, 1971 22(20'). Integument reddish-brown; pronotum with an elliptical discal dark spot; pubescence unicolor, yellowish; lateral margins of pronotum straight from apex of hind angles; pronotum slightly grooved longitudinal medially; hind angles of pronotum backwards; median basal tubercle triangular elongate or indistinct; borders of mesosternal cavity raised at base; median anterior region of metasternum forming a prominence between mesocoxae; free margin of metacoxal plate with small lobe; lateral margins of elytra strongly rounded; third interstice raised near base forming a tubercle (Fig. 13). Fiji and Solomon Islands ...................... Conobajulus Van Zwaluwenberg, 1940 22'. Integument and pubescence coloration different from above; lateral margins of pronotum variable; pronotum not grooved or groove different from above; lateral margins and hind angles of pronotum variable; median anterior region of metasternum not prominent; median basal tubercle of pronotum of other shape; borders of mesosternal cavity not raised at base; median anterior region of metasternum not prominent between mesocoxae; free margin of metacoxal plate without lobe (except Cryptalaus); lateral margins of elytra not rounded; 3rd interstice different from above................................. 23 23(22'). Integument black; pubescence black with small white spots dorsally; 3rd antennomere almost two times longer than 4th; subapical tooth of mandible present; convexity of pronotum moderate from sutures; median basal tubercle of pronotum rounded; borders of mesosternal cavity horizontal (Fig. 6). South Africa, Indonesia (Java)........................... Aliteus Candèze, 1857 23'. Integument and pubescence of varied color; 3rd antennomere shorter than above; subapical tooth of mandible absent; convexity of pronotum variable; median basal tubercle of other shape; borders of mesosternal cavity horizontal or declivous..................................................................... 24 24(23'). Pubescence brownish with dark-brown or black spots; pronotum with two small rounded and elytra with lateral dark pubescent spots; pronotum longer than wide, with transversal median carina near base; lateral margins of pronotum sinuous; convexity of pronotum forming longitudinal median ridge; prosternal channel present; interstices convex; apex of elytra rounded with sutural spine (Fig. 24). Niger, Sierra Leone, Ivory Coast, Ghana, Togo, Cameroon, Central African Republic, Democratic Republic of Congo, Gabon, Angola, Uganda, Tanzania...................................... Neocalais Girard, 1971 24'. Pubescence different from above; length of pronotum variable, without transversal median carina near base; lateral margins of pronotum variable; convexity of pronotum different from above; prosternal channel present or absent; interstices flat or convex; apices of elytra different from above.......................................................................................... 25 25(24'). Integument dark-brown; pubescence with several tonalities of brown; dark pubescence forming two rounded spots on pronotum and a lateral median large spot each elytron; medianterior margin of pronotum sinuous; median basal tubercle of pronotum elongate; notosternal sutures sinuous; metasternal median suture furrowed near base; third interstice raised near base forming a longitudinal ridge; apex of elytra notched with lateroapical and sutural spines; last ventrite of male with truncate apex (Fig. 9). Australia....................................................... Austrocalais Neboiss, 1967 25'. Integument and pubescence different from above; medianterior margin of pronotum of other shape; median basal tubercle of pronotum elongate or transverse; notosternal sutures straight; metasternal median suture furrowed or not near base; 3rd interstice different from above; apex of elytra different from above; last ventrite of male with rounded apex............................. 26 26(25'). Body narrow; integument from reddish- to dark-brown; pubescence brownish of varied tonalities, making regular or irregular patterns; pronotum with 0-4 rounded or elliptical discal darker spots; 3rd antennomere of male transverse with spiniform appendix; metasternal median suture furrowed near base; free margin of metacoxal plate with small lobe; third interstice raised near base forming a dentiform tubercle; apex of elytra notched with lateral angle rounded and sutural spine present (Fig. 15). Australia.............................. ................................................................... Cryptalaus Neboiss, 1967 26'. Body wide and arched or almost parallel; integument and pubescence color different from above; 3rd antennomere of male triangular; metasternal median suture not furrowed near base; free margin of metacoxal plate straight; 3rd interstice different from above; apex of elytra different from above.......................................................................................... 27 27(26'). Integument from reddish- to dark-brown; pubescence black with rusty yellowish-brown setae forming irregular patches; pronotum with longitudinal elliptical groove near base; hind angles of pronotum carinate; borders of mesosternal cavity declivous; metasternal median suture with transversal anterior carina; elytra with apex truncate with lateral spine (Fig. 10). India, Senegal, Guinea, Liberia, Ivory Coast, Ghana, Benin, Cameroon, Gabon, Congo, Democratic Republic of Congo, Tanzania, Angola, South Africa........ ........................................................................ Calais Castelnau, 1836 27'. Integument and pubescence different from above; pronotum not grooved or grooved of other shape; median basal tubercle of pronotum of other shape; hind angles of pronotum not carinate; borders of mesosternal cavity horizontal; metasternal median suture without carina; apex of elytra different from above.................................................................................... 28 28(27'). Integument black and/or reddish; pubescence scale-like, unicolor, white or black dorsally and white ventrally; antennae strongly serrate or pectinate from 4th antennomere; lateral margins of pronotum straight, with hind angles divergent; pronotum strongly convex longitudinal medially and grooved laterally; median basal tubercle of pronotum elongate; medioanterior margin of pronotum raised forming two teeth; prosternal channel present; scutellum subquadrangular; interstices alternate; apex of elytra conjointly rounded (Figs. 32, 33). Ecuador and Bolivia ............................. Propalaus gen. nov. 28'. Integument and pubescence coloration different from above; antennae serrate or pectinate from 3rd antennomere; lateral margins of pronotum variable; pronotum with convexity variable; prosternal channel absent; scutellum of other shape; interstices variable; apex of elytra different from above........................................................................................... 29 29(28'). Pubescence scale-like unicolor, ochrish; fore angles of frons strongly elevate forming a tooth-like; lateral margins of pronotum straight and forming a ridge; convexity of pronotum stronger at middle; posterior margin of pronotum prominent and notched at middle; medioanterior margin of pronotum prominent at middle; meso- metasternal suture present; borders of mesosternal cavity declivous; apex of last ventrite of female rounded; sutural spine of elytra present (Fig. 12). Mexico, Guatemala, Honduras, Costa Rica, Nicaragua, Panama, Colombia, Venezuela, Trinidad and Tobago (Trinidad), French Guiana, Brazil, Ecuador, Peru ..................................... ................................................................ Chalcolepis Candèze, 1857 29'. Pubescence scale-like, uni- or bicolor, different from above; if yellowish-ochre, with lateral ferrugineous bands on pronotum and elytra; fore angles of frons not raised; lateral margins of pronotum slightly sinuous and not forming ridge; convexity of pronotum moderate and gradual; medioanterior margin of pronotum straight or slightly sinuous; meso- metasternal suture absent or obsolete; borders of mesosternal cavity horizontal; apex of last ventrite of female truncate with fringe of spatulate setae; apex of elytra variable (Fig. 31). United States of America, Mexico, Guatemala, Belize, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Cuba, Jamaica, Virgin Islands, Haiti, Dominican Republic, Puerto Rico, Trinidad and Tobago, Venezuela, Colombia, Guyana, Suriname, French Guiana, Brazil, Ecuador, Peru, Bolivia, Chile, Paraguay, Argentina, Uruguay....................................................... ......................................................... Chalcolepidius Eschscholtz, 1829

ACKNOWLEDGEMENTS

To Peterson Lásaro Lopes (Museu de Zoologia, Universidade de São Paulo-MZUSP) by helping with the NDE, TNT and Winclada programs and by discussions on some characters. To Marcelo Duarte (MZUSP) by incentive and valuable suggestions; Paul Johnson (South Dakota State University) by suggestions; Airton de Almeida Cruz (MZUSP) by scanning the diapositives concerning Figs. 31-33; three referees by construtive comments and suggestions. To CNPq for grant.

Recebido em: 12.12.2007

Aceito em: 25.08.2008

Publicado em: 30.09.2008

APPENDIX

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Appendix 1

Appendix 2

Publication Dates

• Publication in this collection
  08 Oct 2008
• Date of issue
  2008

History

• Received
  12 Dec 2007
• Accepted
  25 Aug 2008