Zootaxa, Division of the genus Nothopleurus ...

Zootaxa 2643: 1–44 (2010) www.mapress.com / zootaxa/

ISSN 1175-5326 (print edition)

Article

Copyright © 2010 · Magnolia Press

ZOOTAXA ISSN 1175-5334 (online edition)

Division of the genus Nothopleurus Lacordaire, 1869 (Coleoptera: Cerambycidae: Prioninae) ANTONIO SANTOS-SILVA1, IAN P. SWIFT2 & EUGENIO H. NEARNS3 1

Museu de Zoologia, Universidade de São Paulo. Caixa Postal 42594, 04299-970 São Paulo, São Paulo, Brazil. E-mail: [email protected] 2 California State Collection of Arthropods, 3294 Meadowview Road, Sacramento, CA 95832-1448, USA. E-mail: [email protected] 3 Department of Biology, Museum of Southwestern Biology, University of New Mexico, Albuquerque, NM 87131-0001, USA. E-mail: [email protected]

Table of contents Abstract ............................................................................................................................................................................... 2 Introduction / History .......................................................................................................................................................... 2 Material ............................................................................................................................................................................... 5 Olethrius macrothorax (Montrouzier, 1861) ............................................................................................................... 5 Olethrius cariosicollis (Fairmaire, 1877) .................................................................................................................... 7 Mallodon arabicum Buquet, 1843 ............................................................................................................................... 7 Nothopleurus Lacordaire, 1869 ........................................................................................................................................... 8 Key to the species of Nothopleurus .................................................................................................................................. 10 Nothopleurus subsulcatus (Dalman, 1823)................................................................................................................ 11 Nothopleurus lobigenis Bates, 1884 .......................................................................................................................... 12 Nothopleurus castaneus (Casey, 1924)...................................................................................................................... 15 Nothopleurus madericus (Skiles, 1978)..................................................................................................................... 19 Hovorodon gen. nov. ......................................................................................................................................................... 21 Key to the species of Hovorodon gen. nov. ...................................................................................................................... 22 Hovorodon maxillosum (Drury, 1773) ....................................................................................................................... 23 Hovorodon bituberculatum (Palisot de Beauvois, 1805)............................................................................................ 27 Hovorodon subcancellatum (Thomson, 1867) .......................................................................................................... 28 Hovorodon santacruzensis (Hovore & Santos-Silva, 2004) ...................................................................................... 37 Conclusions ....................................................................................................................................................................... 38 Acknowledgements ........................................................................................................................................................... 39 References ......................................................................................................................................................................... 39

Accepted by Q. Wang: 14 Sep. 2010; published: 13 Oct. 2010

1

Abstract The genus Nothopleurus Lacordaire is reviewed and generic limits redefined. Two species are transferred from Nothopleurus to Olethrius Lacordaire and the following new combinations are proposed: Olethrius cariosicollis (Fairmaire), Olethrius macrothorax (Montrouzier). Nothopleurus arabicus (Buquet) is transferred from Nothopleurus and returned to its original genus as Mallodon arabicum. Mallodon castaneum (Casey) is revalidated in Nothopleurus. Hovorodon gen. nov. is proposed and Cerambyx maxillosus Drury is designated as the type species. Four species are transferred from Nothopleurus to the new genus: Nothopleurus bituberculatus (Palisot de Beauvois), Nothopleurus maxillosus (Drury), Nothopleurus santacruzensis Hovore & Santos-Silva, and Nothopleurus subcancellatus (Thomson). The species of Nothopleurus and of the new genus are redescribed and illustrated. A new country record is recorded for Nothopleurus subsulcatus (Dalman) (Bahamas) and new state records are recorded for N. lobigenis Bates (Mexico: Guerrero, Nayarit). Mallodon hornebecki Chevrolat is considered a valid described species and synonymous with Prionus bituberculatus (= Nothopleurus bituberculatus). In addition, keys are provided to the four species of Nothopleurus as well as the four species of Hovorodon gen. nov. Key words: Macrotomini, new genus, revalidation, taxonomy

Introduction / History Drury (1773) described Cerambyx maxillosus from Barbuda (British West Indies), the oldest species assignable to Nothopleurus. This species was then assigned to Prionus by Fabricius (1775), Prionus (Mallodon) by Drury & Westwood (1837), Mallodon by Thomson (1861), and to Stenodontes (Nothopleurus) by Lameere (1902). Cerambyx maxillosus was described in 1770 (volume I), but it was not until 1773 (volume II) that Drury associated the name with the Linnaean system, following the index to the figures of the first volume. However, references of this species are usually listed as “Drury, 1773: 86, pl. 38, fig.3” (e.g. Monné 2006). Although the year is correct, because the name appeared in 1773 (ICZN 1957: Opinion 474), the page, plate, and figure are from volume 1 (published in 1770). It has also been noted that the index is not paginated (Drury & Westwood 1837; Gemminger & Harold 1872; Gahan 1895; Lameere 1902; mainly Hayek, 1985: 151), however, this is not the case. The index is paginated as “Mm” which corresponds to page “133”. At the time, it was common to identify each page with a number, and for each four pages to include additional identification of the page as a letter (Aa, Bb, Cc, etc). Hayek (1985: 147) pointed out that the name of C. maxillosus appeared in the page 132 (index), but we do not find any page between the page 130 (which is numbered) and the index to the first volume. Thus, C. maxillosus would have been described in Drury, 1773: 131. However, page 129 also has the letters “Ll”, which would indicate the page with the letters “Mm” is the page 133. Palisot de Beauvois (1805) described Prionus bituberculatus from Santo Domingo, Dominican Republic, [Santo Domingo at this time referred to the island of Hispaniola which could have included Haiti or Dominican Republic] which was included in Stenodontes (Nothopleurus) by Lameere (op.cit.), as was Prionus subsulcatus Dalman, 1823, described from Honduras, as well as Mallodon arabicum Buquet, 1843, from the coast of Arabia. White (1853) described Mallodon gnatho from Honduras which was later assigned to Nothopleurus by Waterhouse (1874), and subsequently synonymized under Stenodontes (Nothopleurus) subsulcatus by Lameere (1902). LeConte (1858a) described Mallodon gnatho from Mexico (Sonora). Following LeConte’s description, Gemminger (1872) renamed this species Mallodon mandibulare, to avoid homonymy with Mallodon gnatho White, 1853. Crotch (1873), again renamed Mallodon gnatho LeConte as Nothopleurus dentiger. Although Crotch (op.cit.) had attributed the authorship of the name of the group of species to LeConte, there is nothing in that work that would permit such a conclusion: “The present list is intended only as a means of facilitating exchanges among coleopterists, and hence must not be held as having any authority; synonyms are omitted, except where necessary to connect this work with Dr. Leconte's List from which it has been prepared. Numerous changes have been made in the synonymy, mostly based on comparisons made by Dr. Leconte in

2

· Zootaxa 2643 © 2010 Magnolia Press

SANTOS-SILVA ET AL.

Europe, and already indicated in various papers by himself and Dr. Horn; others on personal examination of the species by myself, and for these I am alone responsible”. This new name later appeared in Horn in Leng (1884) as synonymous of Mallodon mandibularis Gemminger, but also attributed to LeConte in Crotch (op.cit.), and not to Crotch. Curiously, the authorship of the name of the group of species, dentiger, is assigned to Leng (op.cit.) in some works (e.g. Linsley 1962; Monné & Hovore 2006). Mallodon dentiger is not mentioned in some catalogues (e.g. Monné 2006). Montrouzier (1861) described Mallodon macrothorax from New Caledonia, which was later assigned to Olethrius Thomson, 1861 by Lacordaire (1869). Lameere (1903a) affirmed that this species was equal to Opheltes auriculatus Thomson, 1864 from New Caledonia and New Hebrides (Vanuatu), the type-species of Opheltes Thomson, 1864. In the same work, Lameere (op.cit.: 130) synonymyzed Nothopleurus under Opheltes (by priority), and considered Olethrius as a subgenus of Stenodontes Audinet-Serville, 1832. This nomenclatural act was better formalized in Lameere (op.cit.: 139), where Nothopleurus already appears formally in synonymy with Opheltes. Following this action, Lameere (1912) then excluded Olethrius from Stenodontes, considering it a distinct genus. However, Mallodon macrothorax remained as Stenodontes (Nothopleurus) in Lameere (1913, 1919), as well as Opheltes auriculatus being synonymous with Montrouzier’s species. Chevrolat (1862) described Mallodon carptor from Cuba, as well as a second species from Saint Thomas (US Virgin Islands): M. hornebecki. The latter species was not recorded by Thomson (1864, 1867), but was synonymyzed by Gahan (1895) (as M. hornbecki (sic)) under M. carptor. Lameere (1902) recorded M. hornebecki (“sans description”) as synonymous with Stenodontes (Mallodon) bituberculatus, but this name is not included in Lameere (1913), and was mistakenly considered nomen nudum by later authors. This treatment likely had its origin in Gemminger & Harold (1872): “Hornbecki (sic) Chevrol. Ann. Fr. 1862, p. 273. (indscrpt.)”. However, in our opinion, there is no doubt that Mallodon hornebecki Chevrolat, 1862 is a validly described species, as Chevrolat (op.cit.) listed characters (ICZN 1999: Article 12.1) and recorded the type-locality: “Serville (Ann. de la Soc. Ent. de Fr. t. I, p. 176, 1831) ayant pris pour type de son genre Prionus maxillosus F., je ne vois que trios espèces des Antilles, que renferme ma collection, qui doivent y être comprises. Les palpes maxillaires, chez ces insects, ayant le double de longueur des labiaux, et les quatre articles qui les composent étant filiformes allongées, avec le dernier sillonné et terminé en pointe obtuse, ces caractères ne peuvent convenir au grand nombre d’espèces de l’Amérique équinoxiale, qu’on y a joints: de plus, la forme robuste des mandibules, celle du prosternum et du sternum, feront donner un nouveau nom générique à ces insectes…Notre espèce se distinguera de suite de Mallodon maxillosum F., qui n’offre que deux dents internes vers le sommet des mandibules…La troisième espèce est inèdite et originaire d’île SaintThomas (M. Hornebecki nob.)”. From this description it seems clear that Chevrolat (op.cit.), when describing M. carptor, mentioned several characters of M. maxillosum, M. carptor, and M. hornebecki, in addition to pointing out the necessity of a new genus for the continental species. Lacordaire (1869) wrote: “M. Chevrolat (Ann. d. l. Soc. entom. 1862, p. 273), s’appuyant sur quelques differences dans la forme et la longueur des palpes, a proposé également de ne laisser dans le genre que le maxillosus et trois autres espèces des Antilles qu’il ne nomme pas. Mais la différence en question est si légère qu’a peine a-t-elle une valeur de section”. In fact, Chevrolat (op.cit.) mentioned three species (M. carptor, M. maxillosum, and M. hornebecki), and not four as asserted by Lacordaire (op.cit.). In addition, Chevrolat (op.cit.) named these species, contrary to Lacordaire (op.cit.). Chevrolat’s text is confusing, however, it is possible to understand that he was saying that the three species from Antilles, M. maxillosum, M. carptor and M. hornebecki, belong to Mallodon, and that the continental species of Mallodon belonged to a new genus. It is important to note that although a locality alone associated to a name is not an “indication” (ICZN 1999: Article 12.2), in the present case, the name and the locality are associated to the characters shared by the three species studied by Chevrolat (op.cit.). Later, Lameere (1902) placed M. carptor in synonymy with Stenodontes (Nothopleurus) bituberculatus. Thomson (1867) described Mallodon subcancellatum from Brazil, which was synonymyzed, hesitantly, by Lameere (1902) under Stenodontes (Nothopleurus) bituberculatus. This is the same status that Lameere (1903b: 16) maintained for M. subcancellatum: “bituberculatus Beauv., Stenodontes que, toujours dans le même travail, Thomson décrit probablement aussi sous le nom de Mallodon subcancellatum”. In Lameere’s

DIVISION OF THE GENUS NOTHOPLEURUS

Zootaxa 2643 © 2010 Magnolia Press ·

3

catalogue (1913), M. subcancellatum appears in synonymy with Mallodon (Mallodon) spinibarbis (Linnaeus, 1758), without any explanation in this or earlier works. Later, Lameere (1915) revalidated M. subcancellatum as Stenodontes (Nothopleurus) subcancellatus. Lacordaire (1869) erected the genus Nothopleurus for N. ebeninus Lacordaire, 1869 which was designated as a subgenus of Stenodontes by Lameere (1902). Lacordaire (op.cit.) included Olethrius, with four species in his “Remphanides” and divided it in two groups: the first with O. tyrannus and O. scabripennis, in which the labrum is at least as long as wide, in the form of an acute triangle, and abruptly bent down in its apex; the second with O. edwardsii and O. macrothorax, in which the labrum is transverse, subvertical, concave in front, and rounded at the distal margin. Opheltes and Nothopleurus were assigned by Lacordaire (op.cit.) among his “Mallodontides”. Lacordaire (op. cit.) included in Opheltes, O. auriculatus and O. obesus Thomson, 1864. The latter group, according to Lacordaire (op.cit.), does not likely belong in the genus. Lameere (1902) confirmed this when he synonymyzed O. obesus under Stenodontes (Stenodontes) damicornis (Linnaeus, 1771). Fairmaire (1877) described Opheltes cariosicollis from Fiji, and Lameere (1902) assigned it to Stenodontes (Nothopleurus). Nothopleurus lobigenis Bates, 1884, from Mexico (Oaxaca), a second species originally described in Nothopleurus, was considered synonymous with Mallodon gnatho LeConte (= M. mandibularis Gemminger; = M. dentiger) by Lameere (1902), who argued: “Le nom de Bates doit être préféré à celui de Gemminger, parce qu’il y a un Prionus mandibularis Fab. qui est synonyme du Stenodontes exsertus Oliv.”. Lameere (1912: 129) revalidated Nothopleurus, because Opheltes was a homonym of Opheltes Holmgreen (1858), a valid generic name in Hymenoptera. Casey (1912) described Mallodon baroni from Mexico (Guerrero) that was subsequently placed in Nothopleurus by Fragoso & Monné (1995) without further elaboration. Casey (op.cit.), ignoring Lameere (1902), considered Nothopleurus a genus distinct from Stenodontes: “This genus is adopted by Bates, and apparently with ample justification, though rejected by LeConte. Three species are cited by the former author as composing the genus so far as known, of which mandibularis Harold, (= gnatho ║Lec. and dentiger Cr., catalogue name) now stands in our lists under the genus Mallodon; the other species are the type, named ebeninus by Lacordaire, and gnatho White”. Lameere (1919) synonymyzed M. baroni under Stenodontes (Mallodon) spinibarbis (Linnaeus, 1758). Santos-Silva & Martins (2005) placed M. baroni in synonymy with Mallodon dasystomus dasystomus (Say, 1824). According to Lameere (1919), Opheltes is synonymous with Stenodontes (Nothopleurus), which included: S. (N.) arabicus, S. (N.) macrothorax (= Opheltes auriculatus), S. (N.) cariosicollis, S. (N.) subcancellatus, S. (N.) maxillosus, S. (N.) bituberculatus (= Mallodon carptor), S. (N.) subsulcatus (= Mallodon gnatho White and Nothopleurus ebeninus) and S. (N.) lobigenis (= Mallodon gnatho LeConte and Mallodon mandibularis Gemminger). Lameere (1920) transferred Stenodontes (Nothopleurus) cariosicollis and S. (N.) macrothorax to Olethrius and considered the latter genus distinct from Stenodontes: “Un examen approfondi de ce rare Insecte [Olethrius macrothorax], connu également de Kanala à la Nouvelle-Calédonie, m’a permis de me convaincre qu’il devait, de méme que l’espèce cariosicollis FAIRM. des îles Fidji, être retiré des Stenodontes du sous-genre Nothopleurus: ces Longicornes sont des Olethrius, dont ils ont tous les caractères, y compris les épisternums métathoraciques rétrécis em courbe em avant”. However, this nomenclatural act remained unrecognized; thus, both species are listed as belonging to Nothopleurus (e.g. Dillon & Dillon 1952; Fragoso & Monné 1995). Mallodon castaneum Casey, 1924 was described from Mexico (Baja California) and synonymized under Stenodontes (Mallodon) lobigenis by Linsley (1942). Linsley (1957) placed Nothopleurus in synonymy with Mallodon, and maintained it as subgenus of Stenodontes. Quentin & Villiers (1975) disagreed with Linsley (op.cit.), reestablishing Cerambyx spinibarbis Linnaeus, 1758 as the type species of Mallodon, and in so doing, considered Nothopleurus distinct from Mallodon (see Fragoso & Monné 1995).

4


SANTOS-SILVA ET AL.

Hayashi (1961) considered Nothopleurus a distinct genus, but erroneously mentioned Mallodon arabicus as the type species. Skiles (1978) described Stenodontes (Mallodon) madericus from the United States (Arizona) which was assigned to Nothopleurus by Fragoso & Monné (1995). Villiers (1980) reaffirmed the revalidation of Nothopleurus: “Comme nous l’avons démontré (QUENTIN ET VILLIERS , 1975, Faune de Madagascar, 40: 29, c’est par erreur que divers auteurs ont consideré Nothopleurus comme um synonyme de Mallodon S ERVILLE ”. In that same work, they wrote: “genre comprenant quatre espèces d’Amérique centrale et des Antilles. Les autres espèces citées, d’Arabie et des îles du Pacifique appartiennent à um autre genre”. Apparently, Villiers (op.cit.) was also unaware of the transfer by Lameere (1920) of the two species from the Pacific islands. The last two species described in Nothopleurus are: N. santacruzensis Hovore & Santos-Silva, 2004, from Bolivia; and N. komiyai Santos-Silva & Martins, 2005, from Brazil (Bahia). We agree with the exclusion of Mallodon macrothorax and Opheltes cariosicollis established by Lameere (1920), but we do not agree with the inclusion of those species in Olethrius (see below). Mallodon arabicum is also a species that does not belong in Nothopleurus, herein considered an exclusively New World genus. Four of the New World species now assigned to Nothopleurus are therefore transferred to a new genus.

Material The collection acronyms used in this study are as follows: ACMT AMNH BMNH CASC CMSP EMEC FSCA HMUG IRSN MCZN MNHN MNRJ MZSP NHRS MNKM USNM

American Coleoptera Museum (James Wappes), Texas, USA. American Museum of Natural History, New York, USA. The Natural History Museum, London, United Kingdom. California Academy of Sciences, San Francisco, USA. Museum of Stettin, Stettin, Poland. Essig Museum of Entomology, University of California, Berkeley, USA. Florida State Collection of Arthropods, Florida, United States. Hunterian Museum (Zoology), Glasgow University, Glasgow, United Kingdom. Institute Royal des Sciences Naturelles de Belgique, Brussels, Belgium. Museum of Comparative Zoology, Harvard University, Cambridge, USA. Muséum National d’Histoire Naturelle, Paris, France. Museu Nacional do Rio de Janeiro, Rio de Janeiro, Brazil. Museu de Zoologia, Universidade de São Paulo, São Paulo, Brazil. Naturhistoriska Riksmuseet, Stockholm, Sweden. Museo de Historia Natural, Noel Kempff Mercado, Universidad Autónoma “Gabriel René Moreno”, Santa Cruz de la Sierra, Bolivia. National Museum of Natural History, Washington, D.C., USA.

Olethrius macrothorax (Montrouzier, 1861) (Figs. 13, 14) Mallodon macrothorax Montrouzier, 1861: 285. Stenodontes (Nothopleurus) macrothorax; Lameere, 1913: 13 (cat.); 1919: 33. Stenodontes (Opheltes) macrothorax; Lameere, 1903a: 129, 130, 139. Olethrius macrothorax; Lacordaire, 1869: 117; Lameere, 1920: 137. Opheltes auriculatus Thomson, 1861: 306; Lacordaire, 1869: 128; Thomson, 1878: 5 (types); Lameere, 1903a: 129 (syn.). Opheltes macrothorax; Fauvel, 1906: 42. Nothopleurus macrothorax; Hayashi, 1961: 12; Fragoso & Monné, 1995: 219; Komiya & Lorenc, 2006: 4 (checklist). DIVISION OF THE GENUS NOTHOPLEURUS


5

We believe that the inclusion of O. macrothorax (Montrouzier, 1861) in Nothopleurus by Fragoso & Monné (1995) was incorrect, and did not take into account the transfer made by Lameere (1920) from Stenodontes (Nothopleurus) to Olethrius. In fact, the transfer by Lameere (op. cit.) remained unknown to all the authors consulted by us (pers. comm.). Komiya & Lorenc (2006) for example, mention this species in Nothopleurus. Thomson (1861) noted on Olethrius: “Antennae ante oculos insertae, apud ♂ quartam partem posteriorem, et apud ♀ dimidium, attingentes, 11-articulate, art. 1º longissimo, caeteris singulis longiore, 2º– 3º conjunctim 1º aequale, apud ♂ robustioribus; art. 4º precedente multum breviore, 4–11 subaequalibus, gradatim attenuatis. Mandibulae subverticales, apud ♂ magis elongatae et magis robuste, intus excavate, et valde pluridentatae”. This generic description does not permit the inclusion of Mallodon macrothorax in this genus. In males of this species the scape is distinctly short (not surpassing the posterior edge of eyes), antennomeres III–IV together are much longer than the scape, and antennomere III is subequal to IV. Lacordaire (1869), in the redescription of Olethrius wrote that the scape is longer than antennomere III and that this is one-third longer than IV. The inclusion of Mallodon macrothorax in Olethrius, appears to be based on the incorrect identification of this species, as these characters do not apply to Montrouzier’s species. Although the type specimens of M. macrothorax are unavailable for study, we believe that the specimens we examined are of this species based on the type locality, and their close match to the original description and redescription of Lameere (1903a). There is another indication that this species does not possess the characters of Olethrius: Montrouzier (1861) while describing Mallodon edwardsii, immediately before M. macrothorax, wrote in a footnote that this species belongs to Olethrius, whereas in the previous species he did not record such a note. Olethrius macrothorax (Figs. 13, 14), therefore, does not belong to the genus Nothopleurus; the former having the labrum sub-horizontal, distinctly visible in dorsal view, not depressed at the medial region in both sexes (in Nothopleurus it is vertical, not visible or nearly so in dorsal view, and distinctly depressed at the medial region); mandibles of males wider and with dorsal carina low (narrower and with dorsal carina elevated in Nothopleurus); elytral punctation distinct in both sexes (almost smooth in Nothopleurus or, at most, fine and sparse); lateral margins of pronotum crenulated (spined in Nothopleurus); anterior legs proportionally longer, primarily in males (proportionally shorter in Nothopleurus); tibiae without brush of hairs on ventral face in both sexes (with brush in Nothopleurus). Olethrius macrothorax can be included in Hovorodon gen. nov. by similar diagnoses between Nothopleurus, except by the lateral margins of pronotum which are crenulate in both genera, and by the short galea, reaching the apex of the first maxillary palpomere (long in Hovorodon gen. nov. reaching or almost reaching apex of the second maxillary palpomere). In Tavakilian & Chevillotte (1999), this species is recorded as Opheltes macrothorax. If in fact this species is assignable to Olethrius, it will have to be transferred to another genus and, necessarily that genus will have to be Opheltes, if the synonymy established by Lameere between Mallodon macrothorax and Opheltes auriculatus is correct. However, Opheltes will then require a new name, as it will become a junior homonym. We have opted not to take this action without examining the types of M. macrothorax and Opheltes auriculatus. Thus, we provisionally maintain this species in the genus assigned by Lameere (1920), out of which it was never formally transferred. Since M. macrothorax remains incorrectly assigned to Nothopleurus in many works (e.g. Hayashi 1961; Fragoso & Monné 1995), Opheltes should appear as a synonym of this species, however, this is not the case (e.g. Monné 2006). Linsley (1962) was one of the few authors that included Opheltes as synonym of Nothopleurus (mentioned as Stenodontes (Mallodon)). However, due to transfer of M. macrothorax to Olethrius, Opheltes is now a synonym of that genus. Material examined. NEW CALEDONIA, Lifou: 2 males, 1 female [without date and name of collector] (IRSN).

6


SANTOS-SILVA ET AL.

Olethrius cariosicollis (Fairmaire, 1877) (Fig. 15) Opheltes cariosicollis Fairmaire, 1877: 167; 1879: 111; 1881: 470; Lameere, 1883: 6. Stenodontes (Nothopleurus) cariosicollis; Lameere, 1902: 96; 1903a: 130; 1913: 13 (cat.); 1919: 33. Olethrius cariosicollis; Lameere, 1920: 137 (comb. nov.). Nothopleurus cariosicollis; Dillon & Dillon, 1952: 4; Fragoso & Monné, 1995: 219.

Along with Olethrius macrothorax, O. cariosicollis (Fairmaire, 1877) appears incorrectly assigned to Nothopleurus (e.g. Dillon & Dillon 1952; Fragoso & Monné 1995). Likewise, this species does not belong to Olethrius or Opheltes The main characters that prevent the assignment of O. cariosicollis (Fig. 15) to Olethrius are: scape short, reaching at most the posterior edge of eyes; length of antennomere III subequal to IV; prothorax strongly retracted laterally between the posterior and lateral angles, mainly in females; pronotum strongly punctate throughout in both sexes, without impunctate facets. In Olethrius the scape is long, distinctly surpassing the posterior edge of eyes in both sexes, antennomere III is longer than IV (clearer in males), the prothorax is not retracted laterally between the posterior and lateral angles (mainly in males), and the pronotum has impunctate facets in both sexes. The inclusion in Opheltes as established in the original description, or in Nothopleurus or Hovorodon gen. nov. is also precluded, primarily by the shape of the prothorax, the sculpture of the pronotum, and by the presence of short setae on the pronotum and scutellum. In Opheltes, the form and sculpture of the prothorax is very similar to that of Olethrius in both sexes, and of the species assigned to Hovorodon gen. nov. In Nothopleurus the pronotum has a large, smooth, shining area on the disc. In addition, Opheltes and Nothopleurus have the pronotum and scutellum glabrous. As an additional note, the page citation for this species has been cited incorrectly by almost all authors that referenced this species (including Fairmaire, 1879, 1881): instead of page 180, the correct page is 167. Material examined. Female, without labels (IRSN).

Mallodon arabicum Buquet, 1843 (Figs. 12, 55, 56) Mallodon arabicum Buquet, 1843: 330; White, 1853: 46; Thomson, 1867: 98; Lacordaire, 1869: 126; Thomson, 1878: 17 (types); Waterhouse, 1881: 478; Gahan, 1903: 284; Delahaye & Tavakilian, 2009: 39. Mallodon arabicus; Neumann et al., 2004: 138. Stenodontes (Nothopleurus) arabicus; Lameere, 1902: 95; 1913: 13 (cat.); 1919: 33; Gilmour, 1956: 14. Nothopleurus arabicus; Fragoso & Monné, 1995: 219.

We believe that Villiers (1980) was correct when he noted that Mallodon arabicum Buquet, 1843 did not belong to Nothopleurus. We have provisionally maintained it in Mallodon, following the citation of Neumann et al. (2004). Delahaye & Tavakilian (2009) wrote: “Les espèces se répartissent géographiquement entre les régions néotropicale et afrotropicale après réhabilitation de Mallodon arabicum Buquet, 1843, dans son genre originel par NEUMAN et al. (2004)”. However, Neumann et al. (op.cit.) only mention the species in the genus Mallodon, following what they presented in the “Entomologentagung” of 2003 (Bockkäferfauna von Sokotra (Yemen)), when again they only mention Mallodon arabicus, without any elaboration on the assignment of the genus. We could not find any author between Lameere (1919) and Fragoso & Monné (1995), and Neumann et al. (op.cit.) that formally excluded Mallodon arabicum from Nothopleurus. Mallodon arabicum (Fig. 12) differs from the species of Nothopleurus primarily by the hypostomal carina not notably elevated in males; prothorax of male with sexual punctation (fine and densely punctate on the entire ventral and dorsal surface, except laterally on pronotum); pronotal disc with impunctate and shining facets; lateral margins of pronotum of males crenulated; anterior angles of prothorax wide and distinctly projected forward; lateral angles of prothorax of males situated at the same level as the posterior angles;



7

metepisterna moderately narrow and not concave at inner margin; and lobes of tarsomere III (Figs. 55, 56) dorsally flat at apical third. In Nothopleurus the hypostomal carina of males is notably elevated, the prothorax lacks sexual punctation (sexual dimorphism reduced); the pronotal disc lacks impunctate facets in contrast with the remaining surface; the lateral margins of the pronotum with spines; the anterior angles of prothorax are in general, strongly acute (spined); the lateral angles of prothorax of males are placed distinctly before the posterior angles (not in the same level); the metepisterna are narrow and distinctly concave at inner margin; and the lobes of tarsomere III are convex at apical third. It differs from Hovorodon gen. nov. and Mallodon by the head of males, which is somewhat elongate behind the eyes (anterior angles of prothorax not reaching posterior edge of eyes); the elytra more distinctly punctate; the setation of tibiae more conspicuous; and the form of the lobes of tarsomere III. In Hovorodon gen. nov. and Mallodon the head of males is not elongate behind eyes (anterior angles of prothorax reaching or almost reaching the posterior edge of eyes), although some specimens of M. downesii Hope, 1843 have the head slightly elongate behind eyes; elytral punctation finer (in general, micropunctate); the setation of the tibiae is less conspicuous; and the lobes of tarsomere III are convex (Figs. 53, 54). It also differs from Mallodon by the metepisterna being somewhat narrower, particularly in comparison with Mallodon downesii, which is the closest occurring species of Mallodon. Material examined. YEMEN, Socotra Islands: male, [no date of collection], O. Simony col. (IRSN).

Nothopleurus Lacordaire, 1869 Nothopleurus Lacordaire, 1869: 125; Bates, 1879: 8; Lameere, 1912: 129 (revalidation by homonymy); Casey, 1912: 222, 224 (part); Hayashi, 1961: 10, 12 (part); Villiers, 1980: 140 (part); Fragoso & Monné, 1995: 219 (part); Monné, 1995: 12 (cat.; part); Chemsak, 1996: 79; Monné, 2002: 16 (cat.; host; part); Monné & Hovore, 2005: 14 (checklist); 2006: 13 (checklist); Monné, 2006: 54 (cat.; part) Stenodontes (Nothopleurus); Lameere, 1902: 94 (part); 1913: 13 (cat.; part); 1919: 33 (part); Blackwelder, 1946: 552 (checklist; part). Stenodontes (Mallodon); Linsley, 1962: 21 (part); Chemsak & Linsley, 1982: 2 (checklist; part); Chemsak et al., 1992: 15 (checklist; part); Monné & Giesbert, 1994: 6 (checklist; part). Stenodontes (Opheltes); Lameere, 1903a: 130 (syn.; part). Mallodon; Audinet-Serville, 1832: 176; Thomson, 1864: 307 (not Lepeletier & Audinet-Serville, 1830). Type-species. Nothopleurus ebeninus Lacordaire, 1869 (monotypy).

Redescription. Medium (ca. 25.0 mm) to large (ca. 55.0 mm) size, variable intraspecifically. Integument shining, brown to dark-brown. Male. Head large (Figs. 21, 23, 25, 26), without mandibles, ranging from shorter to longer than prothorax; elongated or slightly elongated behind eyes. Longitudinal dorsal furrow of head well marked from base to near occiput, situated in deep depression between antennal tubercles (occasionally to posterior level of upper ocular lobes). Dorsal region of head with or without almost smooth area between eyes; punctation coarse, dense and anastomosed laterally; setation short, sparse, more distinct near eyes. Lateral area behind eyes coarsely and confluently punctate near apex of upper ocular lobes, and partially rugose to wholly rugose in remaining area. Antennal tubercles large, distinct, rounded or subconical at apex, somewhat vertical or projected laterally. Clypeus long or short, rugose or densely punctate, distinctly elevated laterally or sub-flat; setation moderately long and dense (moreso laterally). Labrum sub-triangular, vertical or sub-vertical; central area depressed; setation long and dense. Eyes large, elongate; distance between upper ocular lobes more than two times width of single lobe; distance between lower ocular lobes more than three times width of lobe. Ocular carina narrow, distinct or nearly so. Hypostomal area rugose, uniformly depressed from gula to mentum (Fig. 25), or abruptly and strongly depressed near mentum (Fig. 26); setation sparse to moderately dense (near mentum). Hypostomal carina elevated (Fig. 25) to very elevated (Fig. 21). Maxillary palps not notably long; palpomere III longer than or equal to IV; palpomere IV slightly securiform. Apex of labial palps attaining maxillary palpomere IV or nearly so. Galea long, densely setose, attaining apex of maxillary

8


SANTOS-SILVA ET AL.

palpomere II or nearly so. Mandible variable intra- and interspecifically; dorsal carina elevated to strongly elevated; inner face with moderately long and dense setae. Antennae filiform, attaining basal one-third of elytra; scape not attaining posterior edge of eyes; antennomere III slightly longer than IV; antennomeres X–XI wholly striated. Prothorax transverse. Pronotum sub-smooth on disc, with area of moderately fine punctures at center; laterally rugose-punctate; anterior margin with slight emargination medially; anterior angles from distinctly projected forward to not projected; lateral angles placed at basal one-third, very distinct, spined or sub-spined; posterior angles variable; lateral margins spined (rarely crenulated). Prosternum coarsely and densely punctate laterally, gradually becoming finer and more sparsely towards middle; central region tumid to very tumid. Apex of prosternal process wide, rounded or sub-triangular. Scutellum glabrous, variable interspecifically: uniformly rounded or pentagonal. Elytra glabrous, finely or microscopically punctate; carinae from distinct to absent; apex uniformly rounded or individually rounded; sutural angle with short spine. Metasternum with large area microsculptured and densely punctate (slightly coarse) laterally, setose, distinctly from the sub-triangular central area where punctation clearly finer and setation shorter and disperse, mainly close to metasternal suture. Metepisterna narrow (largest width of middle region 0.15 times length), with inner margin distinctly concave; setation moderately long and very dense. Ventrites I–IV sub-smooth and shining at medial area, microsculptured, opaque and with small granules laterally (sometimes granules present only on ventrites I or I–II); setation sparse, more distinct laterally (sometimes, almost absent). Legs moderately short. Ventral face of tibiae with brush of setae on each side, approximately from middle to apex. Tarsomere V shorter than I–III together. Male genitalia (Figs. 66–68, 70–72): length of lateral lobes from 0.1 to 0.13 times length of tegmen; latero-lower margin of lateral lobes without emargination (Figs. 70–72); dorsal face of lateral lobes with moderately short and dense setation throughout, more dense and longer towards the apex; apex of lateral lobes with long setae, sometimes as long as the lobe; ventral face of lateral lobes with short setae, very disperse at base, longer and more dense on apical one-third; roof longer than lateral lobes to slightly shorter; basal piece longer than lateral lobes and roof together (ca. 0.3 times total length of tegmen); parameres 0.7 to 0.8 times total length of tegmen. Median lobe approximately as long as tegmen; apex of dorsal lobe rounded and with small central emargination; ventral lobe uniformly narrowed towards apex, not projected beyond apex of dorsal lobe to slightly projected; length of apophyses approximately 0.7 times total length of median lobe. Female. Head (Figs. 22, 24, 27) narrower than in male. Distance between lower ocular lobes less than three times width of single lobe. Antennae attaining or just surpassing basal one-third of elytra. Mandibles generally shorter than in males (except when compared with minor males). Hypostomal carina less elevated. Metasternum without lateral area strongly differentiated from central area. Metepisterna slightly concave at inner margin (largest width of middle region 0.22 times of length). Included species. Nothopleurus lobigenis Bates, 1884; N. madericus (Skiles, 1978); N. subsulcatus (Dalman, 1823); N. castaneus (Casey, 1912). Geographical distribution (Fig. 75). Genus with distribution restricted to the Americas. Comments. Nothopleurus differs from Hovorodon gen. nov. in the following manner: prothorax of males without evident sexual dimorphism; pronotum without impunctate and shining facets distinctly contrasting with the remainder of discal surface; hypostomal carina of males in general, notably elevated at all angles; metepisterna of males notably narrow and strongly concave at inner margin (only slightly concave in females). In Hovorodon gen. nov. the prothorax of males shows sexual dimorphism (wholly micropunctate, except laterally on pronotum), the pronotum has impunctate and shining facets clearly contrasting with the remainder of discal surface, the hypostomal carina of males is not notably elevated at all angles, or is elevated only near the mentum, and the metepisternum is not much narrower and not concave at inner margin in both sexes. Thomson (1861) assigned Mallodon to “Mallodonitae”, which included one of the species placed in Nothopleurus. According to Thomson (op.cit.) that group was characterized by: head wide, not elongated posteriorly; antennae short, filiform or sub-moniliform, with the antennomere III short and the next ones larger or equal in length; mandibles horizontal or sub-vertical, robust; prothorax crenulated or multispined laterally; elytra elongated, sub-depressed; mesosternum flat; cylindrical legs; anterior tibiae unarmed (except DIVISION OF THE GENUS NOTHOPLEURUS


9

in Mallodonhoplus Thomson, 1861). Thomson (1864) transferred some genera from “Mallodonitae” to “Macrotomitae”, excluding from the former group the genera having legs with spines. Lacordaire (1869) agreed with Thomson (1864) by maintaining Mallodon in “Mallodontides”, and included Nothopleurus in this group, and recorded on the legs: “toujours inermes” [always unarmed]. Lameere (1902) used “Sténodontines” to designate the group that included Mallodon and Nothopleurus (both as subgenera of Stenodontes), based on the fact that Mallodon was a subgenus of Stenodontes and thus, the name of the group might not be based on it: “Mallodon Downesi Hope est le coryphée d’un groupe auquel la loi de priorité doit faire donner le nom de Sténodontines”. Later, Lameere (1912) divided Macrotomini into eight groups: “Archetypi”, “Basitoxi”, “Cnemoplites”, “Macrotomae”, “Rhaphipodi”, “Xixuthri”, “Titani”, and “Stenodontes”. The last group evidently included the genus Stenodontes, of which Nothopleurus and Mallodon were subgenera. Lameere (1919) characterized the subgroup “Stenodontes” as: body in general large, more or less depressed; eyes not emarginated; antennal tubercles projected and acute; scape elongated and longer than antennomere III; mandibles dorsally with carina; ligulae small and not divided; prothorax wide, explanate and slightly sloped laterally, and with lateral margins crenulated (their sides more parallel in males than in females); prothorax of males with or without sexual punctation; pronotum with shining callosities; legs robust, with or without spines; ventrite V more or less emarginate. This description agrees with that of Thomson (1861), but does not agree with that of Thomson (1864) and Lacordaire (1869), primarily by the inclusion of species in which the legs have spines. Fragoso & Monné (1995), probably following Lameere (1919), used Stenodontina Lameere, 1912 as a subtribe of Macrotomini and assigned the following genera to it: Mallodon, Stenodontes, Nothopleurus, Neomallodon Linsley, 1957, Mallodonhoplus, Physopleurus Lacordaire, 1869, and Olethrius. In Lameere (1919), Opheltes appeared as a synonym of Stenodontes (Nothopleurus). However, as Stenodontina includes Mallodon, the type of “Mallodontitae” and considered a tribe by some authors, there is no doubt that Stenodontina [Stenodontini] is synonymous with Mallodontina [Mallodontini]. It is important to note that Fragoso & Monné (op.cit.) used Stenodontina as a hierarchical group at the same level of “Mallodontitae” (= Mallodontina; = Mallodontini). Contemporary authors (e.g. Adlbauer 1993; Vitali 2008) referring to North American Prioninae, frequently consider Mallodontini as a tribe of Prioninae. This concept is not formally accepted by any of the researchers who work with the New World fauna; most of which assign to Macrotomini all genera that would be included in Mallodontini. This discussion on Mallodontini (= Mallodontina; = Stenodontina) is important since among the species currently assigned to Nothopleurus, some would not even be of the same subtribe (Lameere 1919) or tribe (Villiers 1980). In reference to Lameere (1919), the species of Nothopleurus (as defined herein) would automatically be excluded by the absence of impunctate and shining facets distinctly in contrast with the remaining surface, and by the lateral margins of the prothorax with spines (rarely slightly crenulated). Based on the description of the tribe Mallodontini by Villiers (1980), Nothopleurus would not be included because the prothoracic episternum is not notably wide as in the species of Mallodon and Hovorodon gen. nov. Moreover, Mallodontini sensu Thomson (1861) and Lameere (1919) included genera in which the species (or at least in part) have spines on the legs, while the concept of this group sensu Villiers (op.cit.) might not include these genera, some of which are similar to the species of Mallodon.

Key to the species of Nothopleurus 1. Length of ventrite V less than 0.3 times the basal width. Male .................................................................................. 2 Length of ventrite V greater than 0.4 times the basal width. Female.......................................................................... 5 2(1). Dorsal carina of mandible strongly elevated at basal half, forming a subacute tooth ............................N. subsulcatus Dorsal carina not forming a subacute tooth at basal half ............................................................................................ 3 3(2). Mandibles shorter than head; lateral outer face of mandible strongly tumid; hypostomal area with a plate, parallel to the hypostomal carina ...............................................................................................................................N. madericus Mandibles, usually, longer than head; lateral outer face of mandible not strongly tumid; hypostomal area without plate parallel to the hypostomal carina ....................................................................................................................... .4

10


SANTOS-SILVA ET AL.

4(3). Mandible of major male (usually longer than 40 mm) not notably narrowed at apical fourth (Fig. 36), inner face distinctly oblique, in general, more or less concave; outer face of mandible of minor male distinctly tumid................... ...................................................................................................................................................................N. castaneus Mandible of major male (usually longer than 40 mm) notably narrowed at apical fourth (Fig. 38), inner face almost vertical; outer face of mandible of minor male not notably narrowed .......................................................N. lobigenis 5(1). Dorsal carina of mandible strongly elevated at basal half (Fig.42)............................................................................ .6 Dorsal carina of mandible not strongly elevated at basal half (Figs. 37, 39) .............................................................. 7 6(5). Hypostomal area with a plate parallel to the hypostomal carina ..............................................................N. madericus Hypostomal area without plate parallel to the hypostomal carina..........................................................N. subsulcatus 7(5). Outer face of mandible tumid with the general appearance of dorsal carina, in dorsal view, wide ..........N. castaneus Outer face of mandible more straight with the general appearance of dorsal carina, in dorsal view, not wide ............ ....................................................................................................................................................................N. lobigenis

Nothopleurus subsulcatus (Dalman, 1823) (Figs. 7, 25, 40, 59, 68, 72, 78) Prionus subsulcatus Dalman, 1823: 63. Stenodontes (Nothopleurus) subsulcatus; Lameere, 1902: 100; 1913: 13 (cat.); 1919: 33. Stenodontes (Mallodon) subsulcata; Blackwelder, 1946: 553 (cat.); Chemsak et al., 1992: 15 (cat.); Noguera & Chemsak, 1996: 396 (cat.). Stenodontes (Mallodon) subsulcatus; Monné & Giesbert, 1994: 6 (cat.). Nothopleurus subsulcatus; Fragoso & Monné, 1995: 219; Monné, 1995: 14 (cat.); Turnbow et al., 2003: 4 (checklist); Monné & Hovore, 2005: 14 (checklist); 2006: 13 (checklist); Monné, 2006: 56 (cat.). Mallodon Gnatho White, 1853: 45; Lameere, 1902: 100 (syn.). Mallodon gnatho; Gemminger & Harold, 1872: 2770. Aplagionathus ? Gnatho; Thomson, 1867: 91. Nothopleurus gnatho; Waterhouse, 1874: xviii; Bates, 1879: 8. Nothopleurus ebeninus Lacordaire, 1869: 125; Gemminger & Harold, 1872: 2769; Bates, 1879: 8; Lameere, 1902: 100 (syn.).

Male (Fig. 7). Head, excluding mandibles, as long as or slightly shorter than prothorax, slightly elongated behind eyes. Central area between antennal tubercles and middle of upper ocular lobes with gibbosity on each side of longitudinal dorsal furrow. Dorsal face with coarse punctures, densely and confluent laterally, finer and sparser towards the longitudinal furrow (occasionally with irregular subsmooth area between posterior ocular edge and occiput). Area behind eyes coarsely and confluently punctate close to dorsal face, rough in region between ocular lobes, and rugose toward base of lower ocular lobe. Antennal tubercles rounded to subconical. Clypeus short (approximately 0.25 times width), rugose-punctate with central longitudinal depression, and distinctly elevated laterally. Labrum sub-vertical (only lateral areas visible dorsally). Eyes large; distance between upper ocular lobes from 1.8 to 1.9 times length of scape; distance between lower ocular lobes from 2.0 to 2.3 times length of scape. Ocular carina narrow and distinct. Hypostomal area uniformly depressed from gula to mentum (Fig. 25), generally slightly more marked at central region close to anterior edge; setation sparse. Hypostomal carina elevated (Fig. 25). Maxillary palpomere III as long as IV. Apex of labial palps attaining approximately apex of maxillary palpomere III. Galea not reaching apex of maxillary palpomere II. Mandibles (Fig. 7) as long as the head (minor males) or longer (major males); dorsal carina (Fig. 40) strongly elevated at base, forming tooth, gradually sloped from tooth to level of inner apical tooth, and abruptly sloped (major males), or gradually lowered from apex of tooth to area of abrupt declivity (minor males); inner margin with two large teeth together protracted (minor males) or with large tooth before middle and small tooth near inner apical tooth (major males). Antennae not attaining middle of elytra; scape coarsely punctate. Anterior angles of prothorax not spiniform, projected forward; lateral angles subspinose; posterior angles obtuse or rounded. Elytra finely punctate; elytral carinae distinct. Metepisternum (Fig. 59) notably narrowed and concave at inner margin (width at central region 0.05 times length). Genitalia (Figs. 68, 72).



11

Dimensions in mm (male). Total length (including mandibles), 39.2–45.7; prothoracic length, 5.5–6.6; anterior prothoracic width (between apices of anterior angles), 8.8–10.2; posterior prothoracic width (between apices of posterior angles), 8.9–10.2; humeral width, 10.9–12.0; elytral length, 23.8–26.0. Types, type-localities. Of Prionus subsulcatus: holotype male (photograph examined), from Honduras, deposited at NHRS. Of Mallodon gnatho: holotype male (photograph examined), from Honduras, deposited at BMNH. Of Nothopleurus ebeninus: holotype male, from Mexico (Yucatan), deposited at CMSP. Geographical distribution (Fig. 78). Recorded from Honduras (Dalman 1823), Mexico (Lacordaire 1869) and Guatemala (Monné 1995). Recorded for the first time to Bahamas. Material examined. BAHAMAS, Hope Town: Man-of-War Cay, 2 males, V.9.1986, Feller & Ryan col. (EMEC). Comments. Males of N. subsulcatus are distinguished from those of other species of the genus by the unique large tooth on the mandibular carina or, at least having it strongly elevated at its base.

Nothopleurus lobigenis Bates, 1884 (Figs. 3, 4, 23, 24, 38, 39, 58, 67, 71, 76) Mallodon gnatho LeConte, 1858a: 81; 1858b: 40; 1873: 287; LeConte & Horn, 1883: 272. Mallodon mandibulare Gemminger, 1872: 254; Bates, 1879: 10; Horn in Leng, 1884: 9; Horn, 1895: 337. Mallodon mandibularis; Gemminger & Harold, 1872: 2770. Nothopleurus mandibularis; Bates, 1884: 234; Garnett, 1918: 172 (distr.). Mallodon (Nothopleurus) dentiger Crotch, 1873: 82; Horn in Leng, 1884: 9 (syn.); Bates, 1884: 234 (syn.). Nothopleurus lobigenis Bates, 1884: 235; Fragoso & Monné, 1995: 219 (cat.; part); Chemsak, 1996: 81, pl. 5, figs. 3, 4 (part); Monné, 2002: 16 (cat.; hosts); Noguera et al., 2002: 623 (distr.); Monné, 2004: 7, 13, 20, 27, 61, 66 (cat.; hosts); Santos-Silva & Martins, 2005: fig. 15; Monné & Hovore, 2005: 14 (checklist); 2006: 13 (checklist); Monné, 2006: 55 (cat.). Stenodontes (Nothopleurus) lobigenis; Lameere, 1902: 101; 1913: 14 (cat.); 1919: 33; Linsley, 1934b: 162 (distr.; part); 1935: 69 (distr.); Linsley & Ross, 1940: 76 (biol.); Linsley, 1942: 24 (syn.; distr.; part); Gilmour, 1954: 7, pl. 6, fig. 9. Stenodontes (Mallodon) lobigenis; Linsley, 1957: 4; 1962: 23 (part); Skiles, 1978: 414; Hovore, 1988: 2 (hosts; distr.; part); Chemsak et al., 1992: 15 (distr.); Monné & Giesbert, 1994: 6 (cat.; part); Noguera & Chemsak, 1996: 396 (distr.; part). Stenodontes lobigenis; Brimley, 1938: 209 (distr.); Blackwelder, 1946: 552 (cat.; part); Turnbow & Franklin, 1980: 347 (distr.); Chemsak et al., 1988: 293 (distr.); Chemsak & Noguera, 1993: 57 (distr.); Linsley & Chemsak, 1997: 435 (distr.). Nothopleurus komiyai Santos-Silva & Martins, 2005: 394; Monné & Hovore, 2006: 13 (checklist). Syn. nov.

Male (Fig. 3). Head, excluding mandibles, distinctly longer than prothorax, elongated behind eyes. Central area between antennal tubercles and base of upper ocular lobes, with gibbosity well marked on each side of longitudinal dorsal furrow, followed by punctiform depression. Central area of dorsal face with coarse, sparse punctures at region between gibbosities and middle of region between posterior edge of upper ocular lobe and occiput, and finer and more dense towards latter; laterally strongly vermiculated near upper ocular lobes and coarser and anastomose-punctate towards occiput; setation short and sparse on middle region, longer and more dense laterally. Area behind upper ocular lobe depressed and strongly, longitudinally vermiculated near eye, and rough towards occiput; area behind lower ocular lobe rough and longitudinally rugose close to upper ocular lobe, and longitudinally, strongly vermiculated from middle to base of lobe, and strongly and transversely vermiculated towards occiput. Antennal tubercles rounded, with apex projected laterally. Clypeus short (length ca. 0.27 times width), rugose-punctate, and strongly elevated laterally, forming a wide keel. Labrum sub-vertical (only lateral areas visible dorsally). Eyes proportionally large; distance between upper ocular lobes 1.7 times length of scape; distance between lower ocular lobes 1.8 times length of scape. Ocular carina narrow and distinct. Hypostomal area (Fig. 23) strongly elevated near the gula, and very strongly depressed towards mentum; surface wholly rugose-punctate; setation long, sparse towards gula and more

12


SANTOS-SILVA ET AL.

dense at depressed area. Hypostomal carina (Fig. 23) strongly and gradually elevated from base to apex (major male), with apex curved interiorly. Maxillary palpomere III slightly shorter than IV. Apex of labial palps attaining ca. apex of maxillary palpomere III. Galea not attaining apex of maxillary palpomere II. Mandibles (Fig. 3) longer than head (major male); dorsal carina (Fig. 38) strongly elevated from base to proximity of apical inner tooth, somewhat narrow throughout, and abruptly and obliquely lowered at apex (major male); inner face high, from lower margin to apex of carina at middle region, ca. 0.3 times length of mandible, almost perpendicular to apex of dorsal carina, mainly at middle area, and not notably wide (major male); infero-inner margin with small tooth before middle (major male); outer face more or less flat and oblique at basal half (major male); inner apical tooth distinctly shorter than external. Antennae not reaching middle of elytra; scape dorsally and ventrally coarsely and sparsely punctate, and coarsely and confluently punctate on anterior face; width of scape at apex 0.45 times length. Anterior angles of prothorax with spine just before apex, not projected forward; lateral angles distinctly spinose; posterior angles distinctly projected; lateral margins distinctly spinose. Elytra finely punctate and not slightly rugose; elytral carinae distinct. Metepisternum (Fig. 58) not notably narrowed and concave at inner margin (width at central region 0.12 times length). Genitalia (Figs. 67, 71). Female (Fig. 4). Head, excluding mandibles, slightly longer than prothorax (major female). Sculpture of dorsal face of head, and area behind eyes as in male. Clypeus long (length ca. 0.4 times width), not strongly elevated laterally, but with deep depression at each side. Labrum visible dorsally. Distance between upper ocular lobes 1.7 times length of scape; distance between lower ocular lobes 1.3 times length of scape. Hypostomal area wholly vermiculated, slightly and gradually depressed from gula to mentum, with irregular and transverse depression near anterior edge; anterior edge abruptly vertical; setation as in male. Hypostomal carina (Fig. 24) slightly and gradually elevated from base to apex, more abruptly elevated near apex. Mandibles (Fig. 4) distinctly shorter than head; dorsal carina (Fig. 39) elevated from base to proximity of apical inner tooth, not narrow throughout, and obliquely and gradually lowered to apex; inner face (from lower margin to apex of carina at middle region, distinctly shorter than 0.3 times length of mandible), oblique from inner edge to apex of dorsal carina, mainly at middle area, and notably wide; infero-inner margin without small tooth before middle; outer face somewhat rounded; inner apical tooth distinctly shorter than external. Antennae not reaching basal one-third of elytra; width of scape at apex 0.4 times length; sculpture of scape as in male. Angles and lateral margins of prothorax as in male. Metepisternum slightly concave in inner margin (width at central region 0.25 times length). Variation. Male: Head, excluding mandibles, shorter than prothorax (minor male); central area of dorsal face of head, between antennal tubercles and base of upper ocular lobes, with gibbosity barely marked on each side of longitudinal dorsal furrow (minor male); area behind upper ocular lobe only vermiculate near eye (minor male); area behind lower ocular lobe ranging from rough and longitudinally rugose to simply rugose near upper ocular lobe, and longitudinally and strongly vermiculated to sparsely punctate from middle to base of lobe, or wholly transversely and strongly vermiculated; clypeus from short (major males) to long (minor males) (length from 0.27 to 0.45 times width); clypeus not strongly elevated laterally (not forming a distinct keel); labrum oblique to vertical, partially to wholly visible dorsally; eyes proportionally very large (minor male); distance between upper ocular lobes from 1.3 to 2.0 times length of scape at apex (distinctly larger in major male than in minor male); distance between lower ocular lobes from 1.1 to 2.0 times length of scape at apex (distinctly larger in major male than in minor male); hypostomal area slightly elevated near the gula, and barely depressed towards mentum; surface of hypostomal area from wholly rugose-punctate to distinctly vermiculated on angle and rugose-punctate on depression; hypostomal carina strongly and abruptly elevated at level of depressed region and slightly elevated close to gula (major male), or wholly slightly elevated (minor male); maxillary palpomere III slightly longer than IV; mandibles from shorter to longer than head (distinctly shorter in minor male); dorsal carina of mandible gradually lowered at apical one-third, and not narrow (major male), or not strongly elevated (minor male); inner face very wide, distinctly oblique to apex of dorsal carina, mainly at middle area (minor male), rarely concave or somewhat concave; inner face low (from lower margin to apex of carina at middle region, distinctly shorter than 0.3 times length of mandible); inferoinner margin of mandible without small tooth before middle (major and minor male not present in latter);



13

inner margin of mandibles with one or more teeth together protracted (only in minor male and sometimes in males of medium size); outer face of mandible slightly rounded at basal two-thirds (minor male); antennae surpassing middle of elytra; scape confluently punctate dorsally at basal one-third; scape not confluently punctate at frontal face; width of scape at apex from 0.4 to 0.5 times length; anterior angles of prothorax without spine before apex; anterior angles of prothorax slightly projected forward; lateral angles of prothorax not distinctly spinose; lateral margins of prothorax partially crenulated and with short spines; elytra slightly rugose; width of metepisternum at central region from 0.12 to 0.15 times length. Female: head as long as prothorax (minor female); depression at each side of clypeus not deep; labrum partially visible dorsally; distance between upper ocular lobes from 1.6 to 1.8 times length of scape; distance between lower ocular lobes from 1.2 to 1.4 times length of scape; hypostomal area depressed from middle area to anterior edge; inner apical tooth of mandibles sometimes almost indistinct; width of scape at apex from 0.35 to 0.50 times length; width of metepisternum at central region from 0.20 to 0.25 times length; lateral margins of prothorax somewhat crenulated. Dimensions in mm (male/female). Total length (including mandibles), 22.1–52.8/28.6–52.0; prothoracic length, 3.6–7.3/4.1–7.3; anterior prothoracic width (between apices of anterior angles), 5.3–13.3/6.6–13.0; posterior prothoracic width (between apices of posterior angles), 5.4–13.5/6.3–12.8; humeral width, 6.1–14.3/ 8.2–15.8; elytral length, 14.0–29.3/18.8–33.9. Types, type-localities. Mallodon gnatho (not White, 1853) (Mallodon mandibulare; Mallodon dentiger): holotype male, from Mexico (Sonora) deposited at MCZN. Nothopleurus lobigenis: holotype male, from Mexico (Oaxaca), deposited at MNHN. Nothopleurus komiyai: holotype male, deposited at MZSP. Santos-Silva & Martins (2005) recorded that the type-locality of this species (Brazil, Bahia, Santo Antonio) was doubtful because the species has characters similar to those from Central America. We are therefore excluding the species of the Brazilian fauna, because we believe that the holotype was incorrectly labeled. Geographical distribution (Fig. 76). United States [California (Bates 1884), Arizona (LeConte 1858b), Texas (LeConte 1873), North Carolina (Brimley 1938)], Mexico [Durango (Noguera & Chemsak 1996), Jalisco (Chemsak et al. 1988), Sonora (LeConte 1858), Oaxaca (Bates 1884), Mexico (Linsley 1935), Sinaloa (Bates 1884), Nayarit (new record), Chiapas (new record), Puebla (new record), Morelos (Noguera et al. 2002), Guerrero (new record)]. Material examined. MEXICO, male, [no date indicated], J. Flohr col. (IRSN). Sonora: Alamos, male, VIII.15–20.1958, R. L. Westcott col. (EMEC); Guaymas, male, VIII.31.1968, R. W. Thorp col. (EMEC); (Mira Mar Beach), female, VII.4.1953, Besson col. (EMEC); Navojoa, female, VII.29.1957, R. L. Westcott col. (EMEC); Sonoyta, female, VII.26.1966, J. A. & M. A. Chemsak, E. G. & J. M. Linsley col. (EMEC); 16 km NE Ciudad Obregon, 2 males, VIII.20.1969, J. T. Doyen col. (EMEC). Sinaloa: Choix, 2 males, VII.5.1968, T. A. Sears, R. C. Gardner, C. S. Glaser col. (EMEC); Mazatlan, 2 males, VIII.10–11.1970, J. A. Chemsak col. (EMEC); 6 mi. S Culiacan, 2 males, VIII.06.1964, J. A. Chemsak & J. Powell col. (EMEC); 8 mil. S Elota, female, VII.02.1968, F. D. Parker & L. A. Stange col. (EMEC); 5 mi. N. Mazatlan, male, VII.01.1965, J. A. & M. A. Chemsak, E. G. & J. M. Linsley col. (EMEC); 6 mi. E Villa Union, male, VI.30.1965, J. A. & M. A. Chemsak, E. G. & J. M. Linsley col. (EMEC). Nayarit: Arroyo Santiago (8 mi. NW Jesus Maria), 3 males, 2 females, VII.05.1955, B. Malkin col. (EMEC); Navarrete, 2 males, 2 females, VII.01.1962, A. E. Michelbacher col. (EMEC); male, VII.01.1962, A. E. Michelbacher col. (MZSP); San Blas, female, VI.07.1954, C. Dodson col. (EMEC); male, VI.20.1954, C. Dodson col. (EMEC); Tepic, male, V.20.1958, L. M. Smith col. (EMEC); 2 mi. E San Blas, female, VI.24.1963, W. A. Foster col. (EMEC). Puebla: 45 mi N Acatlan, 1 male, 1 female, VII.30.1963, J. Doyen col. (EMEC). Guerrero: Hwy 200, 41 km NE Ixtapa, male, VII.17–20.1985, J. E. Wappes col. (ACMT). Oaxaca: Matias Romero, male, VIII.14.1963, F. D. Parker & L. A. Stange col. (EMEC); near Cuicatlan, 3 males, VI.28.1972, G. H. Nelson & A. Lau col. (EMEC); 23 mi S Matias Romero, 2 males, IV.5.1962, L. A. Stange col. (EMEC). Chiapas: Tuxtla Gutierrez, 1 male, 1 female, V.03.1959, H. E. Evans col. (EMEC). UNITED STATES, California: Riverside County, Blythe, male, VIII.12.1938, C. Dammers col. (EMEC); male, VII.4.1957, J. W. MacSwain & R. F. Smith col. (EMEC); Coachella, female, VIII.4.1924, F. H. Wymore col. (MZSP); female, VIII.4.1924, F. H. Wymore col.

14


SANTOS-SILVA ET AL.

(MZSP); Palm Beach, N of Earp, male, VIII.25.1962, G. Bateman col. (MZSP); Palm Springs, male, 1932, [no collector indicated] (MZSP); 1 male, 2 females, 1932, [no collector indicated] (EMEC); male, VII.06.1932, [no collector indicated] (EMEC); male, VI.2.1940, [no collector indicated] (EMEC); male, V.24.1940, R. Husbands col. (EMEC). Arizona: Maricopa County, 9 mi N Sunflower, female, VII.15.1964, Jim Haddock col. (EMEC); Pima County, Continental, male, VII.5–10.1973, J. E. Wappes col. (ACMT); Tucson, male, VI.15.1935, O. Bryant col. (EMEC); Organ Pipe Cactus National Museum (Quitobaquito), male, E. Sleeper, W. Agnew, G. Nooman & P. Sullivan col. (EMEC); Yuma County, Yuma, female, VIII.1940, [no collector indicated] (EMEC). Comments. Nothopleurus lobigenis is similar to N. madericus, but differs (both sexes) mainly by the latero-outer face of the mandibles not being notably tumid, and by the absence of a plate on the hypostomal area, parallel to the hypostomal carina.

Nothopleurus castaneus (Casey, 1924) (Figs. 1, 2, 21, 22, 36, 37, 57, 66, 70, 76) Mallodon castaneum Casey, 1924: 225; Linsley, 1942: 24 (syn.). Stenodontes (Nothopleurus) lobigenis; Linsley, 1934a: 59 (distr.).

Male (Fig. 1). Head, excluding mandibles, longer than prothorax, elongated behind eyes. Central area, between antennal tubercles and base of upper ocular lobes with gibbosity well marked on each side of longitudinal dorsal furrow. Central area of dorsal face with coarse, sparse punctures on the region between gibbosities and middle of region between posterior edge of upper ocular lobe and occiput, and finer and more dense towards latter; laterally strongly vermiculated near upper ocular lobes and coarser and anastomosed punctate towards occiput; setation short and sparse on middle region, longer and more dense laterally. Area behind upper ocular lobe depressed and strongly, longitudinally vermiculated near eye, and rough towards occiput; area behind lower ocular lobe less vermiculated than area behind upper ocular lobes. Antennal tubercles rounded, with apex projected laterally. Clypeus short (length ca.0.2 times width), densely punctate, and strongly elevated laterally, forming a wide keel. Labrum sub-vertical (only lateral areas are visible dorsally). Eyes proportionally large; distance between upper ocular lobes 1.9 times length of scape; distance between lower ocular lobes 1.6 times length of scape. Ocular carina narrow and distinct. Hypostomal area (Fig. 21) strongly elevated near gula, and very strongly depressed towards mentum; surface wholly rugosepunctate; setation long, sparse towards gula and more dense at depressed area. Hypostomal carina (Fig. 21) gradually elevated from base to the apex (more distinctly in major specimens), apex curved interiorly. Maxillary palpomere III slightly longer than IV. Apex of labial palps attaining apex of maxillary palpomere III. Galea not attaining apex of maxillary palpomere II. Mandibles (Fig. 1) longer than head (major male); dorsal carina (Fig. 36) strongly elevated from base to proximity of apical inner tooth, somewhat narrow throughout, and not abruptly lowered at apex; inner face high (from lower margin to apex of carina at middle region, ca. 0.3 times length of mandible), oblique to apex of dorsal carina at punctured area (never perpendicular), and notably wide (major and minor specimens); infero-inner margin without small tooth before middle; outer face more or less flat and oblique at basal half (major male); inner apical tooth distinctly shorter than external; apical one-third not notably narrowed (major and minor specimens). Antennae not attaining middle of elytra; scape dorsally and ventrally coarsely and sparsely punctate, and coarsely and confluently punctate on frontal face; width of scape at apex 0.50 times length. Anterior angles of prothorax with spine just before apex, not projected forward; lateral angles distinctly spinose; posterior angles distinctly spinose; lateral margins distinctly spinose. Elytra finely punctate; elytral carinae not distinct. Metepisternum (Fig. 57) not notably narrowed and concave at inner margin (width at central region 0.2 times length). Genitalia (see Figs. 66, 70).



15

FIGURES 1–4. Habitus. 1. Nothopleurus castaneus (Casey, 1924), male; 2. idem, female; 3. Nothopleurus lobigenis Bates, 1884, male; 4. idem, female.

16


SANTOS-SILVA ET AL.

FIGURES 5–8. Habitus. 5. Nothopleurus madericus (Skiles, 1978), paratype male; 6. idem, female; 7. Nothopleurus subsulcatus (Dalman, 1823), male; 8. Hovorodon maxillosum (Drury, 1773), male.



17

Female (Fig. 2). Head, excluding mandibles, slightly longer than prothorax (major female). Sculpturation of dorsal face of head, and area behind eyes as in male. Clypeus long (length ca. 0.3 times width), not strongly elevated laterally, with deep depression at each side. Labrum visible dorsally. Distance between upper ocular lobes 1.8 times length of scape; distance between lower ocular lobes 1.5 times length of scape. Hypostomal area (Fig. 22) wholly vermiculated, slightly and gradually depressed from gula to mentum, with irregular and transverse depression near anterior edge; anterior edge abruptly vertical; setation as in male. Hypostomal carina (Fig. 22) slightly and gradually elevated from base to apex, more abruptly elevated near apex. Mandibles (Fig. 2) distinctly shorter than head; dorsal carina (Fig. 37) elevated from base to proximity of apical inner tooth, wide throughout, and obliquely and gradually lowered to apex; inner face (from inferoinner margin to apex of carina at middle region) distinctly shorter than 0.3 times length of mandible, oblique from inner edge to apex of dorsal carina, mainly at middle area, and notably wide; infero-inner margin without small tooth before middle; outer face somewhat rounded; inner apical tooth distinctly shorter than external. Antennae not reaching basal one-third of elytra; width of scape at apex 0.45 times length; sculpturation of scape as in male. Angles and lateral margins of prothorax as in male. Metepisternum slightly concave in inner margin (width at central region 0.2 times length). Variation. Male: Head, without mandibles, from shorter to slightly longer than prothorax, in major and minor specimens; central area of dorsal face of head, between the antennal tubercles and base of upper ocular lobes, with gibbosity feebly marked on each side of longitudinal dorsal furrow (minor male); central area of dorsal face of head with punctures on region between gibbosities and middle of region between posterior edge of upper ocular lobe and occiput, somewhat dense and from finer to coarser towards latter; dorsal face of head, between upper ocular lobes and occiput, from coarser and anastomosed punctate to strongly rough; area behind upper ocular lobe not or slightly depressed, wholly vermiculated; area behind lower ocular lobe from smooth to rough and slightly rugose close to upper ocular lobe, or wholly transversely vermiculated; clypeus from short (major males) to long (minor males) (length from 0.2 to 0.5 times width); clypeus not strongly elevated laterally (not forming a distinct keel); clypeus somewhat smooth at central area.; labrum almost wholly visible dorsally; distance between upper ocular lobes from 1.8 to 1.9 times length of scape at apex (larger in major male than in minor male); distance between lower ocular lobes from 1.45 to 1.60 times length of scape at apex (larger in major male than in minor male); hypostomal area feebly elevated near gula, and feebly depressed towards mentum; surface of hypostomal area somewhat vermiculated at angle; maxillary palpomere III slightly shorter than IV; mandibles from shorter to longer than head (distinctly shorter in minor male); inner margin of mandible with one or more teeth together protracted (only in minor male and sometimes in males of medium size); outer face of mandible slightly rounded at basal two-thirds (minor male); width of scape at apex from 0.45 to 0.50 times length; anterior angles of prothorax without spine just before apex; anterior angles of prothorax slightly projected anteriorly; width of metepisternum at central region from 0.15 to 0.2 times length. Female: head as long as prothorax (minor female); length of clypeus from 0.30 to 0.45 times width; depression at each side of clypeus not deep; labrum partially visible dorsally; distance between upper ocular lobes from 1.7 to 1.9 times length of scape; distance between lower ocular lobes from 1.45 to 1.55 times length of scape; hypostomal area coarsely and confluently punctate on area near gula; hypostomal area depressed from middle area to anterior edge; inner face of mandible (from lower margin to apex of carina at middle region, not distinctly shorter than 0.3 times length of the mandible); width of scape at apex from 0.45 to 0.55 times length. Dimensions in mm (male/female). Total length (including mandibles), 24.6–48.6/30.3–46.2; prothoracic length, 4.0–6.5/4.0–6.7; anterior prothoracic width (between apices of anterior angles), 5.9–10.5/6.5–9.5; posterior prothoracic width (between apices of posterior angles), 6.0–10.3/7.2–11.8; humeral width, 7.1–13.2/ 8.7–13.0; elytral length, 15.2–27.3/20.0–29.8. Types, type-locality. Two syntypes, male and female, from Mexico, deposited at USNM. The locality of the syntype male is San Felipe, Baja California. The syntype female is from Santa Rosa and, as there are places in Baja California and Baja California Sur with these names, we do not know precisely the correct state of these localities. However, it is likely these localities are in Baja California and Baja California Sur, respectively.

18


SANTOS-SILVA ET AL.

Geographical distribution (Fig. 76). Mexico [Baja California (Casey 1912) and Baja California Sur (first formal record)]. Material examined. MEXICO, Baja California: San Felipe, 2 females, V.18.1963, J. C. Ball col. (EMEC). Baja California Sur: Arroyo ca. 13 miles N. San Ignacio, female, IV.2.1961, Allyn G. Smith col. (EMEC); Cerro Culposo (Cabo Pulmo), male, III.26.1947, [no collector indicated] (EMEC); San José del Cabo, male, IX.11–16.1967, J. Chemsak, A. & M. Michelbacher col. (MZSP); 1.5 miles W San Jose del Cabo, female, VII.17.1968, J. Davidson, J. Bigelow, M. Bentzien, W. Fox, S. Williams, M. Cazier col. (EMEC); 6 miles N San José del Cabo, 3 males, 1 female, IX.15.1967, J. Chemsak, A. & M. Michelbacher col. (EMEC); 1 mile NE San Pedro, 3 males, IX.8.1967, J. Chemsak, A. & M. Michelbacher col. (EMEC); 2 miles NE San Pedro, 10 males, 4 females, IX.19.1967, J. Chemsak, A. & M. Michelbacher col. (EMEC); 3.5 miles NE San Pedro, male, IX.7.1967, J. Chemsak, A. & M. Michelbacher col. (EMEC); 5 miles NE San Pedro, male, IX.1.1967, J. Chemsak, A. & M. Michelbacher col. (EMEC); 7 mi S San Pedro, female, VIII.10.1966, J. Chemsak, J. Doyen, J. Powell col. (EMEC); 3 miles N Santiago, male, VII.16.1957, R.J. & A. Rychman, D. Spencer col. (EMEC); San Lucas, female, VII.20.1968, J. Davidson, J. Bigelow, M. Bentzien, W. Fox, S. Williams, M. Cazier col. (EMEC); 4.5 miles SE La Paz, 2 males, VIII.3.1966, E. & J. Linsley, J. Chemsak, P. D. Hurd col. (EMEC); male, VIII.4.1966, E. & J. Linsley, J. Chemsak, P. D. Hurd col. (EMEC); male, VIII.6.1966, E. & J. Linsley, J. Chemsak, P. D. Hurd col. (EMEC); 5 miles SW La Paz, male, IX.7.1967, J. Chemsak, A. & M. Michelbacher col. (EMEC); female, IX.7.1967, J. Chemsak, A. & M. Michelbacher col. (MZSP); 7 miles SW La Paz, 3 females, VIII.4.1966, E. & J. Linsley, J. Chemsak, P. D. Hurd col. (EMEC); 2 females, VIII.6.1966, J. Chemsak, J. Doyen, J. Powell col. (EMEC); 2 females, IX.6.1967, J. Chemsak, A. & M. Michelbacher col. (EMEC); 20 km S La Paz, male, VIII.26–IX.1.1994, J. E. Wappes col. (ACMT); 10 km NL Los Barriles, female, VIII.30.1994, J. E. Wappes col. (ACMT). Comments. Linsley (1942) synonymyzed Mallodon castaneum with Stenodontes (Nothopleurus) lobigenis, without examining the syntypes. However, he did examine specimens from Baja California and Baja California Sur. We believe that Mallodon castaneum is a different species of Nothopleurus. Nothopleurus castaneus differs from N. lobigenis mainly by the form of the mandibles. The mandible of major males of N. castaneus (Fig. 36) are not notably narrowed, mainly at the apical one-fourth, the inner face is distinctly oblique in general, more or less concave, and the outer-face is, in general, more densely and coarsely punctate. In minor males and in females (Fig. 37) the outer face is distinctly tumid. Major males of N. lobigenis have the mandible notably narrowed (Fig. 38), mainly at the apical one-fourth, the inner face is almost vertical, and the outer-face is in general, finer and sparsely punctate; minor males and females (Fig. 39) do not have the mandible notably tumid at outer face. Another small difference between N. lobigenis and N. castaneus are the punctures of the outer face of mandibles, which are usually coarser and more dense in the latter. Examination of a large series of specimens from Mexico allowed us to conclude that N. komiyai is a junior synonym of N. lobigenis.

Nothopleurus madericus (Skiles, 1978) (Figs. 5, 6, 26, 27, 41, 42, 60, 76) Stenodontes arizonicus; Hovore & Giesbert, 1976: 350; Hovore et al., 1978: 99 (host). Stenodontes (Mallodon) madericus Skiles, 1978: 414; Chemsak et al., 1992: 15; Monné & Giesbert, 1994: 6. Nothopleurus madericus; Fragoso & Monné, 1995: 225; Monné, 1995: 13 (cat.); Chemsak, 1996: 80; Linsley & Chemsak, 1997: 435 (host); Monné, 2002: 16 (cat.; host); 2004: 38 (cat.; host); Monné & Hovore, 2005: 13 (cat.): 2006: 13 (cat.); Monné, 2006: 55 (cat.).

Male (Fig. 5). Head, excluding mandibles, longer than prothorax, elongated behind eyes. Central area, between antennal tubercles and base of upper ocular lobes, with gibbosity well marked on each side of longitudinal dorsal furrow. Central area of dorsal face with coarse, sparse punctures on region between gibbosities and middle region between posterior edge of upper ocular lobe and occiput, and finer and more dense towards latter; laterally strongly coarse and anastomosed near upper ocular lobes and rough towards DIVISION OF THE GENUS NOTHOPLEURUS


19

occiput; setation short and sparse on middle region, longer and more dense laterally. Area behind upper ocular lobe depressed, longitudinally vermiculated near eye, and rough towards occiput; area behind lower ocular lobe rough; setation moderately long and dense. Antennal tubercles rounded, with apex projected laterally. Clypeus long (length equal to approximately 0.3 times width), densely and coarsely punctate, strongly elevated laterally, forming a wide keel. Labrum vertical (only lateral areas are slightly visible dorsally). Eyes proportionally large; distance between upper ocular lobes equal to 1.6 times length of scape; distance between lower ocular lobes equal to 1.3 times length of scape. Ocular carina narrow and distinct between antennal tubercles and posterior edge of upper ocular lobe. Hypostomal area (Fig. 26) strongly elevated near the gula, and very strongly depressed towards mentum; surface wholly rugose-punctate; lateral area near mentum and close to hypostomal carina with a strongly elevated plate at each side, coarsely punctate, and with apex rounded and curved to center of hypostomal area; setation long, sparse towards gula, becoming dense at depressed area and plates. Hypostomal carina (Fig. 26) very low, slightly elevated from base to apex. Maxillary palpomere III longer than IV. Apex of labial palps attaining approximately apex of maxillary palpomere III. Galea not reaching apex of maxillary palpomere II. Mandibles (Fig. 5) longer than head (major male); dorsal carina (Fig. 41) strongly elevated from base to proximity of apical inner tooth, somewhat narrow throughout, and vertically lowered at apex; inner face high (from lower margin to apex of carina at middle region, ca. 0.5 times length of mandible), vertical to apex of dorsal carina at infero-inner margin, and notably wide (major and minor specimens); infero-inner margin with large tooth before middle; outer face tumid; inner apical tooth distinctly shorter than external; apical one-third not notably narrowed (major and minor specimens). Antennae reaching middle of elytra; scape dorsally and ventrally coarsely and sparsely punctate, coarsely and confluently punctate at frontal face; width of scape at apex 0.4 times length. Anterior angles of prothorax with spine just before apex, not projected forward; lateral angles distinctly spinose; posterior angles spinose; lateral margins distinctly spinose. Elytra smooth, usually with the apical fourth finely and sparsely punctate; elytral carinae not distinct. Metepisternum (Fig. 60) distinctly narrowed and concave in inner margin (width at central region ca. 0.12 times length). Female (Fig. 6). Head, excluding mandibles, as long as prothorax. Sculpturation of dorsal face of head and area behind eyes as in male. Clypeus long (length ca. 0.3 times width), not strongly elevated laterally or keel-like, but with deep depression at each side. Labrum visible dorsally. Distance between upper ocular lobes 1.3 times length of scape; distance between lower ocular lobes 1.2 times length of scape. Hypostomal area (Fig. 27) wholly rugose; somewhat elevated near gula, and distinctly depressed towards mentum; anterior edge abruptly vertical; setation as in male. Hypostomal carina (Fig. 27) as in male. Mandibles (Fig. 6) distinctly shorter than head; dorsal carina (Fig. 42) elevated from base to proximity of apical inner tooth, wide throughout, and obliquely lowered at apex; inner face (from infero-inner margin to apex of carina at middle region) ca. 0.5 times length of mandible, concave from inner edge to apex of dorsal carina, and notably wide; infero-inner margin with a plate linking basal tooth with inner apical tooth; outer face tumid. Antennae reaching the basal one-third of elytra; width of scape at apex 0.4 times length; sculpturation of scape as in male. Angles and lateral margins of prothorax as in male. Metepisternum slightly concave at inner margin (width at central region 0.2 times length). Variation. Male: clypeus not punctate at central basal area; clypeus elevated laterally, but not forming wide keel; distance between lower ocular lobes 1.6 times length of scape; ocular carina distinct only on upper ocular lobe; maxillary palpomere III as long as IV; mandibles shorter than head (minor male); anterior angles of prothorax slightly projected forward. Female: head, without mandibles, just shorter than prothorax; distance between upper ocular lobes 1.4 times length of scape; distance between lower ocular lobes 1.4 times length of scape; hypostomal carina indicated; antennae almost reaching the middle of elytra. Dimensions in mm (male/female). Total length (including mandibles), 35.1–41.5/32.2–46.0; prothoracic length, 5.1–5.5/4.6–6.0; anterior prothoracic width (between apices of anterior angles), 8.3–9.5/6.8–10.0; posterior prothoracic width (between apices of posterior angles), 8.5–9.9/7.6–10.4; humeral width, 9.7–10.8/ 9.2–12.4; elytral length, 21.3–24.1/20.7–31.1.

20


SANTOS-SILVA ET AL.

Types, type-locality. Holotype male, from the United States (Arizona), deposited in USNM. Twenty three paratype males and thirty five paratype females, deposited in USNM, CASC, AMNH, EMEC, and private collection of D. D. Skiles, J. S. Cope, E. F. Giesbert, F. T. Hovore, C. E. Langston, A. E. Lewis, D. G. Marqua, and G. C. Walters, Jr. Geographical distribution (Fig. 76). United States [Arizona (Skiles 1978)], Mexico [Chihuahua (Skiles 1978)]. Material examined. UNITED STATES, Arizona: Pima County, Madera Canyon, female, VII.25.1972, B. C. Ratcliffe col. (ACMT); 2 females, VII.24.1978, J. E. Wappes col. (ACMT); Santa Cruz County, Madera Canyon, paratype male, paratype female, VII.11.1957, Stange & Haarding col. (EMEC); Coconino County, Indiana Gardens, 6 miles N Sedona, paratype female, VIII.76.1967, [no collector indicated] col. (EMEC); Superstition Mountains, paratype female, VII [no year indicated]. D. K. Duncan col. (EMEC). MEXICO, Chihuahua: Yepachic, 1 male, 3 females, VII.14.1996, Reyes-Castillo & Edmonds col. (MZSP); 15 miles E Cuauhtemoc (6,600 feet), paratype female, VII.11.1964, J. A. Chemsak & J. Powell col. (EMEC). Comments. Nothopleurus madericus can be easily separated from the other species of the genus by the presence of the plates at the hypostomal area in both sexes (absent in all other species of Nothopleurus). Skiles (1978) affirmed that the mandibles are longer than head. However, the photograph of the holotype (Lingafelter et al. 2010) shows that the mandibles are as long as the head.

Hovorodon gen. nov. Etymology. We are pleased to name this genus in memory of the late Frank T. Hovore for his friendship and many contributions to the study of Cerambycidae. Masculine gender. Type species. Cerambyx maxillosus Drury, 1773. Description. Medium (ca. 24.0 mm) to large (ca. 52.0 mm) size, variable intraspecifically. Integument shining, dark-brown. Male. Head large (Figs. 28, 30, 32, 34), without mandibles, distinctly shorter than prothorax; not elongated behind eyes. Longitudinal dorsal furrow of head well marked from base to near occiput, accommodated in deep depression between antennal tubercles (occasionally to posterior of upper ocular lobes). Dorsal region of head without sub-smooth area between eyes; punctation coarse, dense and anastomosed laterally to entirely; setation short and disperse, more distinct near eyes. Lateral area behind eyes coarse and anastomosed punctate at area closer to apex of upper ocular lobes and rugose in the remainder. Antennal tubercles large, distinct, sub-conic, with apex projected laterally. Clypeus long, with punctation moderately dense, coarse and shallow, not notably elevated laterally; setation moderately long and dense (sometimes more abundant at middle-apical region). Labrum sub-triangular (occasionally with central region strongly elongated), vertical; central area depressed; setation long and dense. Eyes moderately large, elongated; upper ocular lobes placed more than two times width of single lobe, often three; lower ocular lobes placed more than three times width of lobe. Ocular carina narrow, distinct to slightly indistinct. Hypostomal area with variable structure, but always with region closer to mentum distinctly depressed throughout (Figs. 30, 32) or only at central area (Fig. 28, 34); setation disperse to very disperse throughout, moderately dense on depressed area. Hypostomal carina slightly elevated at region closer to mentum (Figs. 28, 30, 32, 34), elevated throughout, obliterated near gula by elevation of hypostomal area, or not elevated. Maxillary palps not notably long; palpomere III as long as, or shorter than IV; palpomere IV slightly securiforme. Apex of labial palps not attaining to surpassing base of maxillary palpomere IV. Galea short, moderately setose, not surpassing the apex of maxillary palpomere I to just surpassing. Mandible variable intra- and interspecifically; dorsal carina elevated to strongly elevated; inner face with moderately long and dense setae. Antennae filiform, just surpassing the middle of elytra; scape variable intraspecifically, generally attaining or just surpassing posterior edge of eyes; antennomere III as long as or just longer than IV; antennomeres X–XI wholly striated.



21

Prothorax transverse. Pronotum finely and very densely punctate (sexual punctation – sexual dimorphism), with impunctate and shining facets on disc (rarely absent in some specimens), rugose-punctate laterally; anterior margin with slight emargination medially; anterior angles wide, distinctly projected forward, or feebly projected; lateral angles placed nearly at same level of posterior angles, distinct, subspinose to rounded or sub-rounded; posterior angles absent or almost so; lateral margins crenulated. Prosternum finely and very abundantly punctate throughout; central region tumid. Apex of prosternal process somewhat narrow, sub-rounded. Scutellum glabrous, variable interspecifically: uniformly rounded to pentagonal. Elytra glabrous or with short and sparse setae on basal one-third, finely to micro-punctate; carinae slightly elevated; apex uniformly rounded (sometimes, sub-truncated); sutural angle with short spine. Metasternum with large setose area, finely, densely punctate and not strongly different from sub-triangular central area, where punctation is distinctly sparser, and setation shorter and less dense, mainly close to the metasternal suture. Metepisterna narrow (largest width of middle region at most 0.25 times length), with inner margin straight to sub-straight; setation moderately long and very dense. Ventrites I–IV sub-smooth and shining at central area, microsculptured, opaque and with granules laterally (occasionally granules absent); setation disperse, more distinct laterally (occasionally nearly absent). Legs moderately short to proportionally longer. Ventral face of tibiae without brush of setae [except in Hovorodon santacruzensis (Hovore & SantosSilva, 2004) that has brush at apical one-fourth of meso- and metatibiae]. Tarsomere V shorter than I–III together. Male genitalia (Figs. 69, 73, 74): length of lateral lobes approximately 0.12 times length of tegmen; latero-lower margin of lateral lobes with emargination (Figs. 73, 74); dorsal face of lateral lobes with moderately short and dense setae throughout to only on apical one-third, more dense and long towards apex; apex of lateral lobes with long setae, occasionally as long as lobe; ventral face of lateral lobes excavated or not, with moderately long and dense setae throughout, longer and dense on apical one-third; roof shorter than lateral lobes, to very shorter; basal piece longer than lateral lobes and roof together (ca. 0.4 times length of tegmen); parameres with 0.7 to 0.8 times total length of tegmen. Median lobe longer or approximately as long as tegmen; apex of dorsal lobe rounded and with small central emargination; ventral lobe uniformly narrowed towards apex, slightly projected beyond apex of dorsal lobe; length of apophyses approximately 0.7 times total length of median lobe. Female. Head (Figs. 29, 31, 33, 35) narrower than in male. Distance between lower ocular lobes slightly less than twice width to 2.5 times width of lobe. Antennae reaching or slightly surpassing the basal one-third of elytra. Mandibles always shorter than in males (except when compared with minor males). Hypostomal carina uniformly sloped from gula to mentum (occasionally somewhat concave). Metasternum as in males. Metepisterna as in males, but width at middle area larger (at most 0.30 times length, but frequently around 0.25 times). Included species. Hovorodon bituberculatum (Palisot de Beauvois, 1805); H. maxillosum (Drury, 1773); H. subcancellatum (Thomson, 1867); H. santacruzensis (Hovore & Santos-Silva, 2004). Geographical distribution (Fig. 75). Genus with distribution restricted to America. Comments. Hovorodon gen. nov. is similar to Mallodon Lepeletier & Audinet-Serville, 1830, from which it differs almost exclusively by the narrow metepisterna (largest width of middle region equal to at most, 0.25 times of length in males and 0.30 in females). In Mallodon the metepisterna are distinctly wider (largest width of middle region equal to at least, 0.30 times the length in both sexes). The concept of Lameere (1902, 1913, 1919), that considered Nothopleurus a subgenus of Stenodontes Audinet-Serville, 1832, which also included Mallodon as subgenus, is not inadmissible. In some species, particularly in females, the metepisternum is not distinctly narrower than in some species of Mallodon.

Key to the species of Hovorodon gen. nov. 1. -

22

Hypostomal area elevated or strongly elevated near of the gula; apex of ventrite V truncated or slightly emarginated. Male .................................................................................................................................................................... 2 Hypostomal area not elevated near of the gula; apex of ventrite V distinctly emarginated. Female.......................... 5


SANTOS-SILVA ET AL.

2(1). 3(2). 4(3).

Fore angle of gena distinctly projected laterally (Fig. 30) ...............................................................H. bituberculatum Fore angle of gena not or slightly projected laterally (Figs. 28, 32, 34) .................................................................... 3 Scape surpassing distinctly the posterior edge of eyes .....................................................................H. santacruzensis Scape, at most, slightly surpassing the posterior edge of eyes................................................................................... .4 Hypostomal area strongly vermiculated and not finely and abundantly punctate on the area elevated near of the gula ................................................................................................................................................................H. maxillosum Hypostomal area, at most, slightly vermiculated on the area elevated near of the gula, and very finely and abundantly punctate throughout .............................................................................................................. H. subcancellatum 5(1). Fore angle of gena distinctly projected laterally (Fig. 31) ...............................................................H. bituberculatum Fore angle of gena not or slightly projected (Fig. 29, 33, 35) laterally....................................................................... 6 6(5). Elytra smooth; humerus, usually, with short and sparse hairs ...............................................................H. maxillosum Elytra usually finely punctate or finely rugose-punctate (rarely almost smooth); humerus without hairs ................. 7 7(6). Scape does not reach the posterior edge of eyes; anterior angles of prothorax somewhat rounded at apex, distinctly projected forward............................................................................................................................. H. subcancellatum Scape reaching the posterior edge of eyes; anterior angles of prothorax somewhat acute at apex, not distinctly projected forward ....................................................................................................................................H. santacruzensis

Hovorodon maxillosum (Drury, 1773) (Figs. 8, 9, 16, 32, 33, 49, 50, 63, 78) Cerambyx maxillosus Drury, 1773: 133; Gmelin, 1790: 1818. Prionus maxillosus; Fabricius, 1775: 163; 1781: 208; 1787: 130; 1792: 249; 1801: 264; Anonymous, 1830: 115 (cat.). Prionus Maxillosus; Schönherr, 1817: 344. Prionus (Mallodon) maxillosus; Drury & Westwood, 1837: 82; Thomson, 1861: 320; Mallodon Maxillosum; Thomson, 1864: 308. Mallodon maxillosum; Thomson, 1867: 103; Gundlach, 1894: 330 (distr.); Gahan, 1895: 83 (distr.); Mallodon maxillosus; Gemminger & Harold, 1872: 2770 (cat.); Fleutiaux & Sallé, 1889: 460 (hosts); Fleutiaux, 1892: 68; Stenodontes (Nothopleurus) maxillosus; Lameere, 1902: 97; 1913: 13 (cat.); 1919: 33; Wolcott, 1951: 332; Duffy, 1960: 58 (Biol.); Gilmour, 1963: 76 (distr.); Peck, 2006: 191 (distr.); Nothopleurus maxillosus; Leng & Mutchler, 1914: 443 (distr.); 1917: 209 (distr.); Wolcott, 1924: 108; 1936: 258; Villiers, 1980: 141; Fragoso & Monné, 1995: 219; Monné, 1995: 13 (cat.); Chalumeau & Touroult, 2005: 45; Santos-Silva & Martins, 2005: fig. 19; Monné & Hovore, 2005: 14 (cat.); 2006: 13 (cat.); Monné, 2006: 55 (cat.); Peck, 2009: 35 (distr.). Stenodontes (Mallodon) maxillosus; Chemsak et al., 1992: 15 (cat.); Monné & Giesbert, 1994: 6 (cat.).

Male (Fig. 8). Central area of dorsal face of head, between the eyes, with punctures distinctly smaller than laterally and towards occiput, somewhat disperse. Area behind upper ocular lobe coarsely punctate, and partially anastomosed; area behind lower ocular lobe rugose. Clypeus slightly elevated laterally; anteriorly distinctly and widely emarginated centrally. Central projection of labrum narrow, acuminated, elongated. Distance between upper ocular lobes equal to 3.7 times the width of scape at apex; distance between lower ocular lobes equal to 4.4 times the width of scape at apex. Hypostomal area (Fig. 32) wholly vermiculated, more distinctly on the area strongly elevated near the gula, and strongly depressed close to the mentum; pilosity moderately short and disperse on elevated area, and distinctly longer and more abundant on depressed area. Hypostomal carina (Fig. 32) elevated throughout, but obliterated near the gula by the elevation of hypostomal area. Apex of gena not projected laterally. Maxillary palpomere III as long as IV. Apex of labial palps not reaching the fourth maxillary palpomere. Galea reaching the apex of the first maxillary palpomere. Mandibles (Fig. 8) distinctly longer than head (major male); dorsal carina (Fig. 49) strongly elevated, narrow throughout, obliquely sloped near the apex; lateroexternal face oblique, somewhat concave, with punctures fine and disperse; pilosity of outer face moderately short and sparse at base, and shorter and sparser in the remaining; pilosity of inner face long and very abundant throughout; infero-inner margin projected in plate at apical half, in which there is a large tooth at apex and the remaining is somewhat irregular; area between the apex of the plate and the apex of inner margin with a large tooth somewhat acute; apex narrow, curved inward, with outer tooth long and acute, and inner tooth short and somewhat rounded. Scape reaching



23

or just surpassing the posterior edge of eyes. Antennomere III longer than IV. Facets of pronotal disc relatively small, subtriangular, placed at anterior half. Lateral angles of pronotum rounded; posterior angles almost null. Elytra with some hairs at basal fifth. Metepisternum (Fig. 63) slightly narrowed and not concave in inner margin (width at central region equal to approximately 0.26 times the length). Female (Fig. 9). Central area of dorsal face of head, between the eyes, coarsely punctate (punctures usually distinctly coarser than in male). Distance between upper ocular lobes equal to 1.8 times the width of a lobe; distance between lower ocular lobes equal to 1.7 times the width of a lobe. Gena as in male. Hypostomal area (Fig. 33) punctate-vermiculate throughout, with transversal furrow, deep, close to the anterior edge. Mandibles (Fig. 50) with long hairs at base of outer face. Lateral angles of pronotum very projected, subspiniform; lateral margins crenulated; disc distinctly punctate, coarser laterally. Elytra with very disperse short hairs at basal fifth. Width of metepisternum equal to 0.30 times the length. Variation. Male: central area of dorsal face of head, between the eyes, with punctures just smaller than laterally and towards occiput, abundant (sometimes partially anastomosed); area behind lower ocular lobe with punctures somewhat fine and sparse on the region close to eye, and rugose towards the base of the head (punctures finer and sparser, and rugosity less conspicuous towards the apex of the eye); distance between upper ocular lobes from 3.6 to 4.0 times the width of scape at apex; distance between lower ocular lobes from 4.1 to 4.4 times the width of scape at apex; hypostomal area not distinctly vermiculated on depressed area; width of metepisternum at central region from 0.26 to 0.28 times the length. Dimensions in mm (male/female). Total length (including mandibles), 53.5–68.5/52.3–53.8; prothoracic length, 9.0–10.2/8.0–8.1; anterior prothoracic width (between the apices of anterior angles), 12.5–15.7/11.0– 11.1; posterior prothoracic width (between the apices of lateral angles), 14.6–18.1/16.2–16.5; humeral width, 14.0–16.4/16.4–16.8; elytral length, 31.5–38.5/35.6–36.1. Types, type-locality. Drury (1773) did not record the sex of the holotype specimen, from Barbuda, but the description and figure from Drury (1770) indicates that the specimen is a male. Unfortunately, the specimens from Drury’s collection are believed to be lost. Ferrer et al. (2004) recorded the following regarding Drury’s collection: “Hayek (1985: 150) lists T. cornutus among Fabrician species described from Drury’s material, but she does not uncover its whereabouts, or that of the majority of Coleoptera types from the Drury Collection. She observes that most of the type material was absent from the sale catalogue prepared upon Drury’s death in 1804, and states that there is ’convincing evidence‘ that 'someone who understood the value of this material acquired it before the collection was offered for sale by public auction‘. Hayek also cites four occasions where Staig (1931, 1940) records possible Fabricius (1775) types described from ’Dom. Drury‘, in the collection of William Hunter…In each case Staig states that Hunter probably acquired the material from Drury…However, Hayek herself refers to the possibility that Drury may have sold part of his collection in or after 1777, when he suffered financial difficulties (and was declared bankrupt), and it seems very possible that part of the type material was acquired, either directly as this time or indirectly later, by Hunter”. There are two specimens of Hovorodon in the HMUG. The specimen with catalogue number GLAHM 139035 is a male of H. bituberculatum. The other specimen (catalogue number GLAHM 139034) (Fig. 16) agrees very well with Drury’s (1770) original description and figure. Dr. Geoff Hancock, Curator of Entomology Department in HMUG, provided the authors with information and photographs of specimens in the Hunter collection. According to Dr. Hancock (pers. comm.): “One photograph [Fig. 65] is of the original drawer which is exactly how they were received here after William Hunter died (1783). The other photograph shows the pair closer up, together with the only label, which they share. The cabinet label gives the information from Fabricius (1781). It is very common in the collection for the insects to be in pairs like this. Often this was because they were thought to be a male and a female. Also it is extremely unusual with these old collections to find any specimen labels on the pin which is a pity. There are none on these specimens’; and: “In some cases we have in Glasgow Drury's specimens of so-called 'missing' types. We also have some Banksian types. Many of these got moved between collections by Fabricius, who was Hunter's curator. His task was to make Hunter's collection as good as possible. So Fabricius took specimens (presumably 'duplicates') from other museums and placed then in Hunter's cabinet and also took some home to Copenhagen or Kiel for his own collection. He was well-known for the redistribution of specimens in this way, as you

24


SANTOS-SILVA ET AL.

probably know. In the case of Drury sometimes Hunter could buy specimens from him. For these reasons it is not necessary to have to explain how Hunter had Drury specimens even though Drury did not die until 1804, many years after Hunter did”. In Anonymous (1784) (it was from these lists that E. Donovan prepared the Drury Sale Catalogue) appears that Drury had nine specimens of Cerambyx maxillosus from Barbuda (near Antigua), collected by W. Killingly in 1767 (with a reference to the illustration 3 of the plate 38). However, when Drury described the species he had a single specimen: “I received it from the island of Barbuda near Antigua, where it was found dead at the foot of a tree”. Drury did not mention the collector or date. Thus, apparently, he acquired the other eight specimens after the description. The Drury sale catalogue (Donovan, 1805) does not mention Cerambyx maxillosus specifically. A number of species of Cerambyx are mentioned by name but then it goes on to say e.g. “16 others or similar”. It is possible that the specimen in the HMUG (GLAHM 139034) is a specimen from Drury’s collection, perhaps even the true holotype of Cerambyx maxillosus. Unfortunately there is no correspondence between Hunter and Drury that might shed any light on this problem (Hunter 2008). Geographical distribution (Fig. 78). Antigua and Barbuda (Barbuda - Drury 1770; Antigua – Chalumeau & Touroult 2005), Cuba (Thomson 1867), Puerto Rico (Gundlach 1894), Guadeloupe (Gahan 1895), Saint Kitts (= Saint St. Christophers) and Nevis (Gahan 1895), Saint Martin (Lameere 1902), Saint Barthélemy (Lameere 1902), Barbados (Duffy 1960), Martinique (Duffy 1960), Dominica (Villiers 1980), Montserrat (Chalumeau & Touroult 2005). Thomson (1867) recorded this species from Cuba, and this country has been listed in catalogues and works as area of distribution of the species. However, Zayas (1957, 1975) did not record H. maxillosum from Cuba. We believe that Thomson (op.cit.) confused H. maxillosum with H. bituberculatum, and thus, the first one needs to be excluded from Cuban fauna, as already pointed out by Gahan (1895) and ignored by all subsequent authors: “In the Catalogue of Gemminger and Harold, Cuba is given as the locality of this species, apparently on the authority of Thomson (Physis i., 2, p. 103), whose citations of localities are not to be always relied upon”. Chalumeau & Touroult (2005) wrote: “Villiers mentione un exemplaire de Martinique [Le Prêcheur, Pointe La Mare (Bonfils)]; il s’agit certainement d’une confusion ave Mallodon spinibarbis – espèce dont Villiers n’avait pas eu connaissance”. We do not know if the specimen mentioned by Villiers (1980) is H. maxillosum. However, according to seen above, Villiers (op.cit.) was not the first to record H. maxillosum from Martinique. Duffy (1960) recorded H. maxillosum to the USA (Florida). After this, the species was not recorded to this country by any author. Probably, the species confused with H. maxillosum by Duffy (op.cit.) is Stenodontes chevrolati Gahan, 1890. Frenzel (1891) recorded this species to Argentina. Without doubt, it was an error of identification. Probably the species is Mallodon spinibarbis (Linnaeus, 1758). Material examined. West Indies, female, [no date indicated], H. A. Ballou col. (USNM). DOMINICA: Clarke Hall Estate (in rotten log), male, 6.VIII.1965, R. M. Anderson col. (USNM). SAINT KITTS & NEVIS: Saint Kitts, 1 male, 1 female (ex-Wickhan collection), 3.V.1904, [name of collector illegible] (USNM). ANTIGUA & BARBUDA: Antigua (Five Islands Peninsula), male, [no date indicated], Raeburn col. (USNM). GUADELOUPE: Les Grands Fonts, female, 27.IV.1979, [no collector indicated] (EMEC). Comments. We believe that the specimens recorded as Stenodontes maxillosus by Redtenbacher (1868) and by Ramos-Elorduy (2006) are not of that species (error of identification). Hovorodon maxillosum is very similar to H. bituberculatum, but can be identified, mainly, by the fore angles of genae without protuberance oriented toward the extending laterally in both sexes (with the protuberance in both sexes of H. bituberculatum).



25

FIGURES 9–12. Habitus. 9. Hovorodon maxillosum (Drury, 1773), female; 10. Hovorodon santacruzensis (Hovore & Santos-Silva, 2004), paratype male; 11. idem, paratype female; 12. Mallodon arabicum Buquet, 1843, male.

26


SANTOS-SILVA ET AL.

Hovorodon bituberculatum (Palisot de Beauvois, 1805) (Figs. 13, 14, 30, 31, 46–48, 62, 74, 78) Prionus bituberculatus Palisot de Beauvois, 1805: 216, pl. 34, fig. 2. Mallodon bituberculatus; Chevrolat, 1852: 649; Gemminger & Harold, 1872: 2770 (cat.). Mallodon bituberculatum; Gahan, 1890: 24. Aplagiognathus bituberculatus; Thomson, 1867: 91. Nothopleurus bituberculatus; Leng & Mutchler, 1914: 443 (dist.); Fragoso & Monné, 1995: 219; Monné, 1995: 13 (cat.); Vitali & Rezbanyai-Reser, 2003: 5; Santos-Silva & Martins, 2005: fig. 18; Chalumeau & Touroult, 2005: 45, 46, fig. 4; Monné & Hovore, 2005: 14 (checklist); 2006: 13 (checklist). Stenodontes (Nothopleurus) bituberculatus; Lameere, 1902: 98; 1913: 13 (cat.); 1919: 33; Gowdey, 1925: 20 (distr.); Russo, 1930: 140 (distr.); Zayas, 1957: 161; 1975: 22. Stenodontes bituberculatus; Wolcott, 1924: 108; Smyth, 1934: 116 (Biology); Wolcott, 1936: 258; 1941: 98; Martorell, 1945: 437 (host); 1945: 208, 245 (host); Ramos, 1946: 41 (distr.). Stenodontes bituberculata: Blackwelder, 1946: 552 (cat.). Stenodontes (Nothopleurus) bitubeculata: Duffy, 1960: 63 (larva). Stenodontes (Mallodon) bituberculata: Chemsak et al. 1992: 15 (cat.). Stenodontes (Mallodon) bituberculatus: Monné & Giesbert, 1994: 6 (cat.). Mallodon carptor Chevrolat, 1862: 273; Thomson, 1867: 91; Gemminger & Harold, 1872: 2770 (cat.); Gahan, 1895: 83. Mallodon hornebecki Chevrolat, 1862: 273; Gahan, 1895: 83 (syn.).

Male (Fig. 13). Central area of dorsal face of head between the eyes with punctures distinctly smaller than laterally and towards occiput, somewhat disperse. Area behind upper ocular lobe somewhat coarsely punctate; area behind lower ocular lobe rugose. Clypeus elevated laterally; anteriorly distinctly and widely emarginated centrally. Central projection of labrum narrow, somewhat acuminated, strongly elongated. Distance between upper ocular lobes 3.5 times width of scape at apex; distance between lower ocular lobes 3.8 times width of scape at apex. Hypostomal area (Fig. 30) wholly vermiculated, strongly elevated near gula, and strongly depressed close to mentum; setation moderately short and disperse on elevated area, and distinctly longer and more abundant on depressed area. Hypostomal carina (Fig. 30) elevated throughout, but obliterated near the gula by elevation of hypostomal area. Apex of gena strongly projected laterally. Maxillary palpomere III as long as IV. Apex of labial palps attaining to slightly not maxillary palpomere IV. Galea not attaining apex of maxillary palpomere I. Mandibles (Fig. 13) distinctly longer than head (major male); dorsal carina (Figs. 46, 47) strongly elevated, narrow throughout, obliquely sloped near apex (major male); latero-external face oblique, somewhat concave, with punctures fine and disperse (coarser and more abundant towards apex and lower margin); setation of outer face moderately long and abundant at base, and shorter and sparse in remaining areas (more conspicuous near lower margin); setation of inner face long and dense throughout; infero-inner margin projected in plate at apical half, where three large teeth present: two near each other at apical one-third of the plate; another at base of plate; apex narrow, curved inward, with outer tooth long and acute, and inner tooth short and somewhat rounded. Scape attaining to almost attaining posterior edge of eyes. Antennomere III slightly longer than IV. Facets of pronotal disc relatively small, subtriangular, placed at anterior half. Lateral angles of pronotum rounded; posterior angles almost absent. Elytra with some setae at basal one-fifth. Metepisternum (Fig. 62) slightly narrowed and not concave on inner margin (width at central region ca. 0.22 times length). Genitalia (Fig. 74). Female (Fig. 14). Central area of dorsal face of head, between eyes, coarsely punctate (punctures distinctly coarser than in male). Distance between upper ocular lobes 2.1 times width of lobe; distance between lower ocular lobes twice width of lobe. Gena as in male. Hypostomal area (Fig. 31) punctatevermiculate throughout, with transverse furrow deep, close to anterior edge. Mandibles (Fig. 48) with long setae at base of outer face. Lateral angles of pronotum very projected, sub-spiniform; lateral margins crenulated; disc almost impunctate. Elytra with very disperse short setae at basal one-fifth. Width of metepisternum 0.30 times length. Variation. Male: central area of dorsal face of head, between eyes, with punctures slightly smaller than laterally and towards occiput, dense (occasionally partially anastomosed); clypeus from slightly to strongly



27

elevated laterally; central projection of labrum not strongly elongated; distance between upper ocular lobes from 3.3 to 3.5 times width of scape at apex; distance between lower ocular lobes from 3.7 to 4.0 times width of scape at apex; hypostomal area not distinctly vermiculated on depressed area; apex of labial palps from not attaining to surpassing base of maxillary palpomere IV; galea not or attaining apex of maxillary palpomere I; mandibles slightly longer than head (minor male); dorsal carina of mandibles with abrupt declivity near apex (almost vertical) (minor male); setation of outer face of mandibles nearly entirely glabrous but with long setae at base; plate of infero-inner margin of mandibles with small and irregular teeth between basal and medial teeth; lateral angles of pronotum sub-spinose to rounded or sub-rounded; posterior angles of pronotum absent; width of metepisternum at central region from 0.22 to 0.25 times length. Female: Distance between upper ocular lobes 2.4 times width of lobe; hypostomal area vermiculate throughout; lateral margins of pronotum with sub-spiniform projections. Dimensions in mm (male/female). Total length (including mandibles), 47.4–55.0/45.5–54.5; prothoracic length, 8.0–9.3/7.3–9.3; anterior prothoracic width (between apices of anterior angles), 11.5–12.5/10.3–11.8; posterior prothoracic width (between apices of lateral angles), 13.3–15.8/14.9–17.7; humeral width, 12.4– 15.4/14.0–17.2; elytral length, 28.9–33.5/30.8–37.9. Types, type-locality. Of Prionus bituberculatus: holotype female, from Dominican Republic (Santo Domingo), not found. Of Mallodon carptor: holotype male, from Cuba, deposited at BMNH. Of Mallodon hornebecki: number of specimens, and sex(es) unknown, from Saint Thomas Island, probably deposited at BMNH. Geographical distribution (Fig. 78). Dominican Republic (Palisot de Beauvois 1805), Cuba (Chevrolat 1852), Jamaica (Gahan 1895), Saint Thomas [Chevrolat (1862), extended to Virgin Islands by (Chalumeau & Touroult 2005)], Puerto Rico (Lameere 1902), Haiti (Blackwelder 1946), and Saint Johns (Chalumeau & Touroult 2005), Guadeloupe (new record). Material examined. PUERTO RICO: Road 385, female, 13.IV.1971, J. Micheli col. (USNM); Mayagüez, female, V.1975, J. Micheli col. (MZSP); Road 385, km 3.1, male, 2.VI.1972, J. Micheli col. (MZSP). CUBA, Las Villas: Sierras de Trinidad, male, V.1986, [no collector indicated] (MZSP). GUADELOUPE: 2 males (exNonfried collection), [no date and collector indicated] (IRSN). Porto Cabello (locality unknown – probably is not Puerto Cabello in Venezuela), female, [no date and collector indicated] (IRSN). Currently, only H. maxillosum is recorded from Guadeloupe [very rarely collected on this island, according to Chalumeau & Touroult (2005)]. The two males examined by us, actually belong to H. bituberculatum, and we do not know if the specimens were incorrectly labeled. Comments. Gahan (1890) noted: “Judging from the figure and description of this species it seems to me that it is the female of Mallodon maxillosum, Drury”. Lameere (1902) wrote: “je pense que le Cerambyx bituberculatus est bien la femelle du Mallodon carptor et non celle du Mallodon maxillosus comme l’a supposé M. Gahan”. However, the description and figure in Palisot de Beauvois (1805) does not permit us to adequately determine the specimen which currently is treated as H. bituberculatum or H. maxillosum. As the latter species is not recorded from Dominican Republic, we believe the specimen is in fact H. bituberculatum. Hovorodon bituberculatum is similar to H. maxillosum, but differs mainly by the form of the apex of gena in both sexes, which is distinctly projected laterally (not projected in H. maxillosum).

Hovorodon subcancellatum (Thomson, 1867) (Figs. 15, 16, 28, 29, 43, 44, 45, 61, 69, 73, 77) Mallodon subcancellatum Thomson, 1867: 102; 1878: 5 (types); Lameere, 1902: 98(syn.). Mallodon subcancellatus; Gemminger & Harold, 1872: 2771 (cat.). Stenodontes (Nothopleurus) subcancellatum Lameere, 1915: 51 (reval.); 1919: 33. Stenodontes (Mallodon) subcancellatus; Monné & Giesbert, 1994: 6 (checklist). Nothopleurus subcancellatus; Fragoso & Monné, 1995: 222; Monné, 1995: 14 (cat.); Santos-Silva & Martins, 2005: fig. 16; Monné & Hovore, 2005: 13 (checklist); 2006: 13 (checklist); Monné, 2006: 56 (cat.).

28


SANTOS-SILVA ET AL.

FIGURES 13–16. Habitus. 13. Hovorodon bituberculatum (Palisot de Beauvois, 1805), male; 14. idem, female; 15. Hovorodon subcancellatum (Thomson, 1867), male; 16. idem, female.



29

Male (Fig. 15). Dorsal face of head wholly coarsely, confluently punctate. Area behind upper ocular lobe coarsely punctate; area behind lower ocular lobe rough. Clypeus slightly elevated laterally; anteriorly distinctly and widely emarginated centrally. Central projection of labrum triangular and slightly projected. Distance between upper ocular lobes 3.5 times width of scape at apex; distance between lower ocular lobes 3.8 times width of scape at apex. Hypostomal area (Fig. 28) tumid, strongly depressed at centro-basal area, wholly micropunctate; area tumid with dense and fine punctures at central area near gula, gradually coarser and confluent towards hypostomal carina and mentum; area depressed vermiculate-punctate; setation short and sparse in tumid area, and slightly longer and more dense on depressed area. Hypostomal carina (Fig. 28) not elevated throughout. Apex of gena slightly projected laterally. Maxillary palpomere III as longer than IV. Apex of labial palps attaining or not maxillary palpomere IV. Galea slightly surpassing apex of maxillary palpomere I. Mandibles (Fig. 15) as long as head; dorsal carina (Figs. 43, 44) strongly elevated, somewhat thickened throughout, obliquely sloped near the apex; latero-outer face oblique, with punctures fine and disperse; setation of outer face moderately long and dense at base, and shorter and almost absent in remaining areas; setation of inner face long and dense throughout; infero-inner margin projected in plate at apical half, three large teeth present; apex narrow, curved inward, with outer tooth long and acute, and inner tooth short. Scape attaining or almost surpassing the posterior edge of eyes. Antennomere III slightly longer than IV. Facets of pronotal disc relatively small, irregular, placed at anterior half, wholly finally punctate, but more shining and less punctate than remaining surface. Anterior angles of pronotum distinctly projected forward; lateral angles distinctly projected; posterior angles almost absent. Elytra wholly glabrous. Metepisternum (Fig. 61) slightly narrowed and not concave at inner margin (width at central region approximately 0.25 times length). Genitalia (Figs. 69, 73). Female (Fig. 16). Central area of dorsal face of head as in male, with similar variation. Area behind upper ocular lobes as in male. Distance between upper ocular lobes 3.3 times width of scape at apex; distance between lower ocular lobes 3.6 times width of scape at apex. Gena as in male. Hypostomal area (Fig. 29) not tumid, punctate-vermiculate throughout, without transverse furrow at anterior edge, uniformly sloped from gula to mentum, sometimes with antero-central area depressed. Mandibles (Fig. 45) with short and sparse setae at base of outer face. Lateral angles of pronotum very projected, sub-spiniform. Elytra as in male. Width of metepisternum 0.30 times length. Variation. Male: central area of dorsal face of head, between eyes, with punctures partially not confluent, and somewhat smaller than punctures close to eyes; punctures of dorsal face of head, between the eyes and occiput, partially not confluent; area behind upper ocular lobe confluently punctate; anterior margin of labrum uniformly rounded; distance between upper ocular lobes from 3.4 to 4.0 times width of scape at apex; distance between lower ocular lobes from 3.6 to 4.4 times width of scape at apex; dorsal carina narrow throughout; setation of outer face of mandible short and sparse at base; facets of pronotal disc sparsely punctate; facets of pronotal disc large; lateral angles of pronotum distinctly rounded; elytra with very short and sparse setae near humeri; metepisternum slightly concave; width of metepisternum at central region from 0.25 to 0.30 times length. Female: area behind upper ocular lobes rough; distance between upper ocular lobes from 3.0 to 3.4 times width of scape at apex; distance between lower ocular lobes from 3.1 to 3.8 times width of scape at apex; apex of genae not projected forward; hypostomal area somewhat depressed. Dimensions in mm (male/female). Total length (including mandibles), 25.0–48.4/36.0–45.4; prothoracic length, 4.5–8.7/5.9–7.8; anterior prothoracic width (between apices of anterior angles), 6.2–12.5/7.5–9.4; posterior prothoracic width (between apices of lateral angles), 7.4–14.5/9.9–13.4; humeral width, 7.7–13.6/ 10.6–13.5; elytral length, 17.0–29.5/25.4–30.3. Types, type-locality. Syntypes males, from Brazil, deposited at MNHN. Thomson (1867) did not specify how many specimens he had, but according to Tavakilian & Chevillotte (1999) there are two syntypes. Geographical distribution (Fig. 77). Brazil [Bahia, Minas Gerais, Rio de Janeiro, Santa Catarina (Fragoso & Monné 1995), São Paulo (Monné 1995), Goiás, Minas Gerais (new record)].

30


SANTOS-SILVA ET AL.

FIGURES 17–20. Habitus. 13. Olethrius macrothorax (Montrouzier, 1861), male; 14. idem, female; 15. Olethrius cariosicollis (Fairmaire, 1877), female; 16.Hovorodon maxillosum (Drury, 1773), probable holotype male (ex-Hunter’s collection, deposited at HMUG).



31

FIGURES 21–32. Head and prothorax, ventral view. 21. Nothopleurus castaneus, male; 22. idem, female; 23. N. lobigenis, male; 24. idem, female; 25. N. subsulcatus, male; 26. N. madericus, paratype male; 27. idem, female; 28. Hovorodon subcancellatum, male; 29. idem, female; 30. Hovorodon bituberculatum, male; 31. idem, female; 32. Hovorodon maxillosum, male.

32


SANTOS-SILVA ET AL.

FIGURES 33–56. 33–35, head and prothorax, ventral view: 33. Hovorodon maxillosum, female; 34. H. santacruzensis, paratype male; 35. idem, paratype female. 36–52, mandibles: 36. Nothopleurus castaneus, male, dorsal view; 37. idem, female, dorsal view; 38. N. lobigenis, male, dorsal view; 39. idem, female, dorsal view; 40. N. subsulcatus, male, lateral view; 41. N. madericus, male, lateral view; 42. idem, female, dorsal view; 43. Hovorodon subcancellatum, male, dorsal view; 44. idem, male, lateral view; 45. idem, female, dorsal view; 46. H. bituberculatum, male, dorsal view; 47. idem, male, lateral view; 48. idem, female, dorsal view; 49. H. maxillosum, male, dorsal view; 50. idem, female, dorsal view; 51. H. santacruzensis, male, dorsal view; 52. idem, female, dorsal view. 53–56, metatarsus: 53. Mallodon downesii Hope, 1843, male, dorsal view; 54. idem, lateral view; 55. M. arabicum, male, dorsal view; 56. idem, male, lateral view.



33

FIGURES 57–64. Metepisternum, male. 57. Nothopleurus castaneus; 58. N. lobigenis; 59. N. subsulcatus; 60. N. madericus; 61. Hovorodon subcancellatum; 62. H. bituberculatum; 63. H. maxillosum; 64. H. santacruzensis.

34


SANTOS-SILVA ET AL.

FIGURE 65. Photograph of the drawer as it was received by the HMUG after the death of William Hunter.

FIGURES 66–74. 66–69. Tegmen, lateral view: 66. Nothopleurus castaneus; 67. N. lobigenis; 68. N. subsulcatus; 69. Hovorodon subcancellatus. 70–74. Lateral lobes, lateral view: 70. Nothopleurus castaneus; 71. N. lobigenis; 72. N. subsulcatus; 73. H. subcancellatum; 74. H. bituberculatum.



35

FIGURES 75–78. Maps. 75. Distribution of Nothopleurus and Hovorodon; 76. Nothopleurus lobigenis (circle), N. castaneus (square), N. madericus (star); 77. Hovorodon subcancellatum (circle), H. santacruzensis (square); 78. H. bituberculatum (square), H. maxillosum (star), Nothopleurus subsulcatus (circle).

Material examined. BRAZIL, Goiás: Vianópolis, 2 males, XI.1931, R. Spitz col. (MZSP). Minas Gerais: Água Suja, male, 5.XI.1919, Stain col. (MZSP). Espirito Santo: Santa Teresa, male, II.1964, C. T. Elias col. (MZSP). São Paulo: Botucatu, female, 2.XII.1971, A. Netto col. (MZSP); male, 14.XI.1972, Maria A. col. (MZSP); male, 29.V.1974, Valemia col. (MZSP); male, 21.XI.1978, M. Barros col. (MZSP); female, 29.XI.1978, Alexandre col. (MZSP); Castilho (left margin Road Paraná), male, X.1964, Expedição Museu de Zoologia col. (MZSP); Guarujá, male, 16.XII.1926, Melzer col. (MZSP); Mongaguá, female, 15.XII.2006, P. L. Soto col. (MZSP); Pindamonhangaba (Eugênio Lefévre; 1200 m), female, 24.I.1963, Expedição Museu Zoologia col. (MZSP); Piracicaba, male, 20.X.1960, K. Reichardt col. (MZSP); Pirassununga, male, [no date indicated], J. Gaspar col. (MZSP). Comments. The general appearance of Hovorodon subcancellatum is very similar to that of the species of Mallodon. It differs mainly by the hypostomal area of males being tumid, and by the narrower metepisterna. As noted earlier, Lameere (1902) hesitantly placed Mallodon subcancellatum in synonymy with Stenodontes (Nothopleurus) bituberculatus, and noted that the specimens in Thomson’s collection were likely

36


SANTOS-SILVA ET AL.

mislabeled. Later, Lameere (1915) revalidated the species: “La collection renferme un exemplaire malle d’un Nothopleurus qui répond complètement à la description du Mallodon subcancellatum J. Thoms. que j’aien eu le tort de paser en synonymie”. However, the specimen examined by Lameere may not have been the same species in Thomson’s collection. According to Lameere (op.cit.), the metepisterna are concave, which does not agree with the specimens from Brazil. Inexplicably, although Lameere (1915) affirmed that the specimen examined by him was from “Amérique méridionale”, he later (Lameere 1919) recorded the distribution of Stenodontes (Nothopleurus) subcancellatus as “Haiti ?”. This species appears incorrectly synonymized with Mallodon spinibarbis in Chemsak et al. (1992).

Hovorodon santacruzensis (Hovore & Santos-Silva, 2004) (Figs. 10, 11, 34, 35, 51, 52, 64, 77) Nothopleurus santacruzensis Hovore & Santos-Silva, 2004: 51; Santos-Silva & Martins, 2005: fig. 17; Monné & Hovore, 2005: 13 (checklist); 2006: 13 (checklist); Wappes et al., 2006: 4 (checklist); Wappes et al., 2009: 1–8 (types)

Male (Fig. 10). Central area of head, between the antennal tubercles and middle of upper ocular lobes, with gibbosity slightly marked on each side of longitudinal dorsal furrow. Punctures of dorsal face distinctly coarser and deeper laterally and towards occiput. Antennal tubercles coarsely punctate at basal half and distinctly finer at apical half, mainly at apex. Length of clypeus approximately 0.35 times width; frontal edge strongly emarginated; surface not strongly sloped in relation to the front. Distance between upper ocular lobes 3.2 times width of scape at apex; distance between lower ocular lobes 3.7 times width of scape at apex. Ocular carina narrow and well marked from base of antennal tubercles to posterior upper ocular edge. Hypostomal area (Fig. 34) distinctly depressed centrally near mentum; surface coarsely vermiculate, wholly, finely punctate; setation long and sparse towards gula and shorter towards mentum. Hypostomal carina (Fig. 34) not elevated. Maxillary palpomere III as long as IV. Apex of labial palps reaching base of maxillary palpomere IV. Galea surpassing the apex of maxillary palpomere I. Mandibles (Fig. 10) approximately 1.2 times longer than head (major male); dorsal carina (Fig. 51) narrow and strongly elevated, obliquely sloped near apex; outer face moderately densely punctate; inner face convex; inner lower margin projected as a plate at apical half, and with some teeth, more distinct at apical third of the plate. Scape surpassing posterior edge of eyes; somewhat coarsely punctate, mainly laterally and dorsally. Anterior angles of prothorax feebly projected forward; posterior angles variable, generally rounded. Elytra glabrous, finely punctate and somewhat finely rugose throughout. Width of metepisterna (Fig. 64) 0.25 times of length. Female (Fig. 11). Head, without mandibles, 0.7 times length of prothorax. Sculpture of dorsal face of head, and area behind eyes as in male. Labrum occasionally longer in male (from 0.3 to 0.4 times width). Distance between upper ocular lobes 2.8 times width of scape at apex; distance between lower ocular lobes 3.2 times width of scape at apex. Mandibles (Fig. 52) almost glabrous on outer face. Antennae attaining basal one-third of elytra; scape not surpassing posterior edge of eyes. Hypostomal area (Fig. 35) without transverse furrow near anterior edge. Width of metepisterna 0.3 times length. Variation. Male: distance between lower ocular lobes from 3.7 to 3.9 times width of scape at apex; hypostomal area not strongly vermiculate throughout; setation of hypostomal area short towards gula and very short and sparse towards mentum; mandible as long as head (minor male); elytra somewhat finely rugose only laterally. Female: head, without mandibles, from 0.7 to 0.8 times length of prothorax; hypostomal area somewhat concave. Dimensions in mm (male/female). Total length (including mandibles), 20.2–42.8/28.9–35.5; prothoracic length, 3.4–6.9/4.6–6.2; posterior prothoracic width (between apices of posterior angles), 6.1–12.0/8.0–10.1; humeral width, 6.0–11.5/8.5–11.1; elytral length, 13.6–24.6/18.9–25.2.



37

Types, type-locality. Holotype male, from Bolivia (Santa Cruz), deposited at MNKM. Forty six paratypes (32 males and 14 females) from the same locality of holotype, deposited at FSCA, EMEC, MZSP, MNRJ, and MCNZ. Hovore & Santos-Silva (2004) mistakenly recorded the holotype as deposited in the USNM collection. Wappes et al. (2009) recorded the holotype as deposited at MNKM. Geographical distribution (Fig. 77). Species recorded only from Bolivia (Santa Cruz) (Hovore & Santos-Silva 2004). Material examined. BOLIVIA, Santa Cruz: Flora & Fauna Hotel, 3.7 km SSE Buena Vista, paratype male, paratype female, XI.5–15.2001, M.C. Thomas & B. K. Dozier col. (MZSP); 4–6 km SSE Buena Vista, paratype male, paratype female, XI.1–8.2002, J. E. Wappes col. (MZSP); female, X.3–8.2004, J. E. Wappes & R. F. Morris col. (MZSP). Comments. Hovorodon santacruzensis resembles H. subcancellatum, but males differ from this species by: mandibles longer and narrower at apical third; dorsal carina of mandibles sub-linear; scape surpassing the posterior edge of eyes; hypostomal area not tumid, vermiculate; anterior prothoracic angles short and feebly projecting anteriorly. In males of H. subcancellatum the mandibles are shorter and slightly narrowed at apical third, the dorsal carina of mandibles is convex at middle, the scape usually does not surpass the posterior edge of eyes, the hypostomal area is tumid, not vermiculated, or very slightly vermiculate antero-medially, and the anterior angles of prothorax are large and distinctly projected forward. Females of H. santacruzensis differ from those of H. subcancellatum by the scape reaching the posterior margin of eye, and by the anterior angles of prothorax being narrowed and feebly or not at all projecting anteriorly. In females of H. subcancellatum the scape does not reach the margin of eye, and the anterior prothoracic angles are relatively wide and strongly projecting.

Conclusions Prioninae is a relatively small subfamily of Cerambycidae (Monné & Hovore 2006). Nevertheless, the taxonomic relationships among the genera are complex. In many cases, the assignment of species into the various genera is extremely artificial. Often, the generic assignments are based on general morphology, not taking into consideration a broader set of characters. Unfortunately, there are cases in which the allocation in genera is completely random. In these instances, even the general shape of the body was used for proper (or at least more logical) assignment. Hovorodon gen. nov. was established to separate it from Nothopleurus species that share exclusive characters which cannot be included in the currently known genera. We believe that Hovorodon and Nothopleurus are monophyletic genera, with a good set of distinctive characters between them, and with other genera of Macrotomini. However, any hypothesis on the monophyly of these genera requires an appropriate phylogenetic analysis and was beyond the scope of this work. Thus, this study provides insights into a future revision of the tribe whose monophyly remains untested. (Santos-Silva & Galileo, 2010). Mallodon is in need a complete revision, due to several problems including incorrect synonymies, incorrect records for certain areas, etc. This is one of the genera in which the species show notable variability, not associated with geographical distribution (this is very common in Prioninae, but particularly evident in Mallodon). As we have noted, the assignment of M. arabicum was provisionally transferred to Mallodon, as it is necessary to examine a largest quantity of specimens. Thus, we believe it is impossible (or at least reckless) to provide a key to the species of Mallodon. In the case of Olethrius, we have no specimens of any species of the genus, so we are unable to provide a useful key. Likewise, the present state of understanding of Macrotomini (and of its genera) does not allow for the construction of a reliable key. However, for the American genera of Macrotomini, Hovorodon can be included in the alternative of couplet “8”, from Santos-Silva & Martins (2005): 8(6). Metepisterna narrow (largest width equal to about 1/4 of the length); inner margin frequently concave .......... .8A 8A(8). Prothorax of males without evident dimorphism; pronotum of males without impunctate and shining facets distinctly contrasting with the remainder of discal surface; metepisterna of males (Fig. 59) notably narrow and

38


SANTOS-SILVA ET AL.

-

-

strongly concave at inner margin, and slightly concave in females ........................ Nothopleurus Lacordaire, 1869 Prothorax of males with evident dimorphism; pronotum of males with impunctate and shining facets clearly contrasting with the remainder of discal surface; metepisterna of males (Fig. 63) not much narrower and not concave at inner margin in both sexes ................................................................................................... Hovorodon gen. nov. Metepisterna wide (largest width equal to about 1/3 of the length); inner margin straight or slightly convex ...... 9

Acknowledgements We greatly appreciate the loan of specimens from Steven W. Lingafelter (USNM), Alain Drumont (IRSN), Cheryl B. Barr (EMEC), James Wappes (ACMT), and Daniel Heffern (Houston, TX). We are also grateful to Sharon Shute (BMNH) for assistance with bibliographic issues and Geoff Hancock (HMUG) for assistance with Drury specimens.

References Adlbauer, K. (1993) Cerambyciden aus Senegambien (Coleoptera: Cerambycidae). Koleopterologische Rundschau, 63, 279–292. Anonymous (1794) A Catalogue of the Exotic Insects in the Collection of Dru Drury. Ca. 200 leaves. Anonymous (1830) Catalogue of the contents of the Museum of Royal College of Surgeons in London. Part IV, fascilus I: Comprehending the first division of the preparations of natural history in spirit. London, Royal College of Surgeons of England, vii + 55 p. Audinet-Serville, J.G. (1832) Nouvelle classification de la famille des longicornes. Annales de la Société Entomologique de France, 1, 118–201. Bates, H.W. (1879) Insecta. Coleoptera, Cerambycidae, Prionidae. Biologia Centrali-Americana, British Museum (Natural History), London, 5, 1–16, pls. I–II. Bates, H.W. (1884) Insecta, Coleoptera. Supplement to Longicornia. Biologia Centrali-Americana, British Museum (Natural History), London, 5: 225–248, pl. XVI. Blackwelder, R.E. (1946) Checklist of the coleopterous insects of Mexico, Central America, the West Indies and South America. Part 4. Bulletin of the United States National Museum, 185(4), 551–763. Brimley, C.S. (1938) The insects of North Carolina, Being a List of the Insects of North Carolina and Their Close Relatives. North Carolina Department of Agriculture, Division Entomology, 560 pp. Buquet, J.B.L. (1843) Description d’une nouvelle espèce de Longicorne de la tribu des Prioniens, appartenant au genre Mallodon de Serville. Revue Zoologique, 1843, 330. Casey, T.L. (1912) III - Studies in the Longicornia of North America. Memoirs on the Coleoptera, 3, 215–376. Casey, T.L. (1924) I - Additions to the known Coleoptera of North America. Memoirs on the Coleoptera, 11, 1–347. Chalumeau, F. & Touroult, J. (2005) Les longicornes des Petites Antilles (Coleoptera, Cerambycidae). Taxonomie, éthologie, biogéographie. Pensoft, Moscow, 241 pp. Chemsak, J.A. (1996) Illustrated Revision of the Cerambycidae of North America. Volume I. Parandrinae, Spondylidinae, Aseminae, Prioninae. Wolfsgarden Books, Burbank, 1: i–x + 150 p., est. I–X. Chemsak, J.A. & Linsley, E.G. (1982) The Longhorned beetles. Checklist of the Cerambycidae and Disteniidae of North America, Central America, and the West Indies (Coleoptera). Plexus, New Jersey, 138 pp. Chemsak, J.A., Linsley, E.G. & Hovore, F.T. (1988) A Concentration Site for Cerambycidae in Jalisco, Mexico (Coleoptera). The Pan-Pacific Entomologist, 64(3), 291–295. Chemsak, J.A., Linsley, E.G. & Noguera, F.A. (1992) Listados Faunisticos de Mexico II. Los Cerambycidae y Disteniidae de Norteamerica, Centroamérica y las Indias Occidentales (Coleoptera). Universidad Nacional Autónoma de Mexico, Instituto de Biologia, México, 204 pp. Chemsak, J.A. & Noguera, F.A. (1993) Annotated checklist of the Cerambycidae of the estacion de Biologia Chamela, Jalisco, Mexico (Coleoptera), with descriptions of new genera and species. Folia Entomologica Mexicana, 89, 55– 102, 3 figs. Chevrolat, L.A.A. (1852) Rappel des coléoptères décrits par Palisot de Beauvois aux genres actuellement adoptés, avec synonymie. Annales de la Société Entomologique de France, (2)10, 629–651. Chevrolat, L.A.A. (1862) Coléoptères de l'Ile de Cuba. Notes, synonymies et descriptions d'espèces nouvelles. Familles des Cérambycides et des Parandrides. Annales de la Société Entomologique de France, (4)2, 245–280. Crotch, G.R. (1873) Check list of the Coleoptera of America, north of México. Naturalists' Agency, Salem, 136 pp. Dalman, J.W. (1823) Analecta entomologica. Holmiae viii + 108 pp., 4 pls.



39

Delahaye, N. & Tavakilian, G.L. (2009) Not sur Mallodon downesii Hope, 1843, et mise en synonymie de M. Plagiatum Thomson, 1867 (Coleoptera, Cerambycidae). Bulletin de la Société entomologique de France, 114(1), 39–45. Dillon, L.S. & Dillon, E.S. (1952) Cerambycidae of the Fiji Islands. Bulletin of Bernice P. Bishop Museum, 206, 1–114. Donovan, E. (1805) (compiler). A catalogue of the most capital assemblage of insects probably ever offered to public sale : consisting of upwards of eleven thousand different specimens, collected from all the countries ... by Mr. Dru Drury... lately deceased, who was upwards of thirty years in forming the above collection : at the same time will be sold the elegant mahogany cabinets, with a few books on Entomology and the copy-right, copper-plates in number one hundred and fifty of the [Drury's] illustrations of natural history ... with a few coloured sets, and a large quantity of letter-press, and odd plates coloured and plain, which will be sold by auction by Messr. King & Lochee ... on Thursday, May 23, 1805 and the two following days at twelve o’clock. 16 pp. Drury, D. (1770) Illustrations of natural history. Wherein are exhibited upwards of two hundred and forty figures of exotic insects, according to their different genera; very few of which have hitherto been figured by any author, being engraved and coloured from nature, with the greatest accuracy, and under the author's own inspection, on fifty copper-plates. With a particular description of each insect: interspersed with remarks and reflections on the nature and properties of many of them. To which is added, a translation into French. D. Drury, London, 1: xxvii + 130 pp., 50 pls. Drury, D. (1773) Illustrations of Natural History, wherein are exhibited upwards of two hundred and twenty figures of exotic insects according to their different genera; very few of which have hitherto been figured by any author, being engraved and coloured from nature, with the greatest accuracy, and under the author’s own inspection, on fifty copper-plates. With a particular description of each insect: interspersed with remarks and reflections on the nature and properties of many of them. D. Drury, London, 2: vii + 90 pp, 50 pls. Drury, D. & Westwood, J.O. (1837) Illustrations of exotic entomology, containing upwards of six hundred and fifty figures and descriptions of foreign insects, interspersed with remarks and reflections on their nature and properties. Henry G. Bohn, London, 1: 1–123, 151 pls. Duffy, E.A.J. (1960) A monograph of the immature stages of Neotropical timber beetles (Cerambycidae). British Museum (Natural History), London, vii + 327, 176 figs & 13 pls. Fabricius, J.C. (1775) Systema entomologiae, sistens insectorum classes, ordines, genera, species, adiectis synonymis, locis, descriptionibus, observationibus. Korte; Flensburgi & Lipsiae, 30 + 832 pp. Fabricius, J.C. (1781) Species insectorum exhibentes eorum differentiae specificas, synonyma auctorum, loca natalia, metamorphosin adiectis observationibus, descriptionibus. Bohn, Hamburgi et Kilonii, 1: viii + 552 pp. Fabricius, J.C. (1787) Mantissa insectorum sistens eorum species nuper detectas adiectis characteribus genericis, differentiis specificis, emendationibus, observationibus. Proft, Hafniae, 1: xx + 348 pp. Fabricius, J.C. (1792) Entomologia systematica emendata et aucta. Secundum classes ordines, genera, species adjectis synonymis, locis, observationibus, descriptionibus. Proft, Hafniae, 1(2): xx + 538 pp. Fabricius, J.C. (1801) Systema eleutheratorum secundum ordines, genera, species: adiectis synonymis, locis, observationibus, descriptionibus. Bibliopoli Academici Novi, Kiliae, 2: 1–687. Fairmaire, L. (1877) Diagnoses de Coléoptères australiens et mélanésiens. Petites Nouvelles Entomologiques, 2(180), 166–167. Fairmaire, L. (1879) Descriptions de coléoptères nouveaux ou peu connus du Musée Godeffroy. Journal des Museum Godeffroy, 5(14), 80–114. Fairmaire, L. (1881) Essai sur les Coléoptères des îles Viti (Fidgi). Suite 1. Annales de la Société Entomologique de France, 6(1), 461–492. Fauvel, A. (1906) Faune Analytique des coléoptères de la Nouvelle-Calédonie. 4e. partie. Revue d’Entomologie, 25, 29– 100. Ferrer, J., Maxwell, V.L. & Geoffrey, H. (2004) Discovery of lost J. C. Fabricius (1775) and A. G. Olivier (1795) types of Coleoptera in the Hunterian Museum, University of Glasgow, Scotland. Annales Zoologici, 54(4), 775–781. Fleutiaux, E. (1892) Petite faune élémentaire des Coléoptères de la Guadeloupe à l'usage des écoles de la Colonie. Imprimerie du Gouvernement, Basse Terre, 87 pp. Fleutiaux, E. & Sallé, A. (1889) Liste des Coléoptères de la Guadeloupe et descriptions d'espèces nouvelles. Annales de la Société Entomologique de France, (6)9, 351–484, pls 7–8. Fragoso, S.A. & Monné, M.A. (1995) Notes on Macrotomini (Coleoptera, Cerambycidae, Prioninae). Revista Brasileira de Biologia, 55(2), 215–225. Frenzel, J. (1891) Uebersicht über eine Coleopterensammlung Von Córdoba in Argentinien. Entomologische Nachrichten, 17, 326–333. Gahan, C.J. (1890) Notes on some West-Indian Longicorn Coleoptera, with Descriptions of new Genera and Species. The Annals and Magazine of Natural History, 6(6)31, 23–34. Gahan, C.J. (1895) On the Longicorn Coleoptera of the West Indian Islands. The Transactions of the Entomological Society of London, 1895, 79–140. Gahan, C.J. (1903) Arthropods insects: Coleoptera. In: Forbes H. O.: The natural history of Sokotra and Abd-el-Kuri. A

40


SANTOS-SILVA ET AL.

monograph of the Islands. Bulletin of the Liverpool Museum, Special Bulletin, 261–292. Garnett, R.T. (1918) An annotated list of the Cerambycidæ of California. (Col.). The Canadian Entomologist, 50(5), 172–177. Gemminger, M. & Harold, E. (1872) Catalogus coleopterorum hucusque descriptorum synonymicus et systematicus. Sumptu E. H. Gummi (G. Beck), Monachii, 9, 2669–2988. Gemminger, M. (1872) Geänderte Namen. Coleopterologische Hefte, 10, 254. Gilmour, E.F. (1954) Notes on a collection of Prioninae (Col., Cerambycidae) from the Institut Royal des Sciences Naturelles de Belgique. Bulletin de l'Institut Royal des Sciences Naturelles de Belgique, 30, 1–48. Gilmour, E.F. (1956) Revision of the “Prioninae” of tropical and South Africa. Longicornia, 3, 1–252. Gilmour, E.F. (1963) Some Caribbean Coleoptera Cerambycidae. Studies on the Fauna of Curaçao & other Caribbean Islands, 18(78), 75–102, 3 pls. Gmelin, J.F. (1790) Caroli a Linné Systema Naturae. Edit. 13. Beer, Lipsiae, 1(4), 1517–2224. Gowdey, C.C. (1925) Catalogus insectorum Jamaicensis. Entomological Bulletin of the Department of Agriculture of Jamaica, 4(1), xiv + 114 pp. Gundlach, J. (1894) Apuntes para la fauna Puerto-Riqueña. Octava parte. Anales de la Sociedad Española de Historia Natural, 2(22)2, 286–344. Hayashi, M. (1961) Cerambycidae from New Caledonia (Col.). Part I. Bulletin of the Osaka Museum of Natural History, 13, 7–65. Hayek, C.M.F. (1985) On the type of material of the species of Coleoptera described from the Drury Collection by D. Drury and J. C. Fabricius with notes on some Coleoptera from the Milne collection preserved in the British Museum (Natural History). Archives of Natural History, 12(1): 143–152. Horn, G.H. (1884) In: Leng C. W. Synopses of Coleoptera (Cerambycidae). Bulletin of the Brooklyn Entomological Society, 7(1), 7–11. Horn, G.H. (1895) The Coleoptera of Baja California. Proceedings of the California Academy of Sciences, 4: 302–449. Hovore, F.T. (1988) Additions to the cerambycid beetle fauna of Baja California, Mexico: records and descriptions (Coleoptera, Cerambycidae). The Wasmann Journal of Biology, 46(1–2), 1–29, 4 figs. Hovore, F.T. & Santos-Silva, A. (2004) Notes and description of Neotropical Macrotomini (Coleoptera, Cerambycidae, Prioninae). Revista Brasileira de Entomologia, 48(1), 49–54. Hunter, W. (2008) The correspondence of Dr. William Hunter, 1718–1783. Edited by Brock, C.H. Pickering & Chatto, London. 800 pp. ICZN (International Commission on Zoological Nomenclature) (1957) Opinion 474. Determination of the dates to be assigned for the purposes of the Law of Priority to the names published in Dru Drury’s Illustrations of Natural History in the period 1770–1782. Opinions and Declarations rendered by the International Commission on Zoological Nomenclature, 16(16), 297–306. ICZN (International Commission on Zoological Nomenclature) (1999) International Code of Zoological Nomenclature. London, xxx + 306 p. Komiya, Z. & Lorenc, J. (2006) Checklist of the Cerambycidae (Subfamily Prioninae) of the world. Part I: Tribes Macrotomini, Platygnathini, Eurypodini. Available from: http://www.cerambycidae.cz (01.07.2009). Lacordaire, J.T. (1869) Histoire Naturelle des Insectes. Genera des Coléoptères, Librairie Encyclopedique de Roret, Paris, 8, 1–552. Lameere, A.A.L. (1883) Liste des Cérambycides décrits postérieurement au catalogue de Munich. Annales de la Société Entomologique de Belgique, 26, 1–78. Lameere, A.A.L. (1902) Révision des Prionides. Quatrième Mémoire – Sténodontines. Mémoires de la Société Entomologique de Belgique, 9, 63–110. Lameere, A.A.L. (1903a) Révision des Prionides. Cinquième mémoire. – Olethrius. Annales de la Société Entomologique de Belgique, 47(3), 129–140. Lameere, A.A.L. (1903b) Révision des Prionides. Septième mémoire. – Macrotomines. Mémoires de la Société Entomologique de Belgique, 11, 1–216. Lameere, A.A.L. (1912) Révision des Prionides .Vingt-deuxième Mémoire. - Addenda et Corrigenda. Mémoires de la Société Entomologique de Belgique, 21, 113–188. Lameere, A.A.L. (1913) Cerambycidae: Prioninae. Coleopterum Catalogus (pars 52). Berlin, W. Junk & S. Schenkling (ed.), 22, 1–108. Lameere, A.A.L. (1915) Note sur les Prioninae du Museum National d'Histoire Naturelle de Paris. Bulletin du Muséum National d'Histoire Naturelle de Paris, 21, 51–63. Lameere, A.A.L. (1919) Famille Cerambycidae: subfam. Prioninae. Coleoptera. In: Genera insectorum. Belgique, P. Wytsman (ed.), (172), 1–189, pls. 1–8. Lameere, A.A.L. (1920) Prioninae nouveaux ou peu connus. Annales de la Société Entomologique de Belgique, 60, 137– 145. LeConte, J.L. (1858a) Description of New Species of Coleoptera, chiefly collected by the United States and Mexican



41

Boundary Commission, under Major W. H. Emory, U.S.A. Proceedings of the Academy of Natural Sciences of Philadelphia, 10, 59–89. LeConte, J.L. (1858b) Catalogue of Coleoptera of the Regions adjacent to the Boundary Line between the United States and Mexico. Journal of the Academy of Natural Sciences of Philadelphia (ser. 2), 4, 9–42. LeConte, J.L. (1873) Classification of the Coleoptera of North America. Prepared for the Smithsonian Institution. Part II. Smithsonian Miscellaneous Collections, 11 (265), 279–348. LeConte, J.L. & Horn, G.H. (1883) Classification of the Coleoptera of North America. Prepared for the Smithsonian Institution. Smithsonian Miscellaneous Collections, 26(507), i–xxvii + 1–567. Leng, C.W. & Mutchler, A.J. (1914) A Preliminary List of the Coleoptera of the West Indies as Recorded to January 1, 1914. Bulletin of the American Museum of Natural History, 33(30), 391–493. Leng, C.W. & Mutchler, A.J. (1917) Supplement to Preliminary List of the Coleoptera of the West Indies. Bulletin of the American Museum of Natural History, 37(5), 191–220. Lingafelter, S.W., Monné, M.A. & Nearns, E.H. (2010) Online Image Database of Cerambycidae Primary Types of the Smithsonian Institution. Available from: http://www.elaphidion.com/ (Accessed on 8/1/2010) Linsley, E.G. (1934a) Studies in the Cerambycidae of Lower California (Coleoptera). The Pan-Pacific Entomologist, 10(2), 59–63. Linsley, E.G. (1934b) Notes and Descriptions of West American Cerambycidae (Coleoptera). Entomological News, 45(6), 161–165. Linsley, E.G. (1935) Studies in the Longicornia of Mexico (Coleoptera: Cerambycidae). Transactions of the American Entomological Society, 61, 67–102, 1 fig., 1 pl. Linsley, E.G. (1942) Contribution toward a knowledge of the insect fauna of Lower California Nº 2 Coleoptera: Cerambycidae. Proceedings of the California Academy of Sciences, (fourth series), 24(2), 21–96. Linsley, E.G. (1957) Descriptive and Synonymical Notes on Some North American Cerambycidae (Coleoptera). American Museum Novitates, 1828, 1–21. Linsley, E.G. (1962) The Cerambycidae of North America. Part II. Taxonomy and Classification of the Parandrinae, Prioninae, Spondylinae & Aseminae. University of California. Publications in Entomology, 19, 1–102. Linsley, E.G. & Ross, E.S. (1940) Records of some Coleoptera from the San Jacinto Mountains, California. The PanPacific Entomologist, 16(2), 75–76. Linsley, E.G. & Chemsak, J.A. (1997) The Cerambycidae of North America, Part VIII: Bibliography, Index, and Host Plant Index. University of California Press, Berkeley, 117, i–ix + 1–534. Martorell, L.F. (1945) A survey of the forest insects of Puerto Rico. The Journal of Agriculture of the University of Puerto Rico, 29(3), 69–354. Martorell, L.F. (1945) A survey of the forest insects of Puerto Rico. The Journal of Agriculture of the University of Puerto Rico, 29(4), 355–608, 18 figs. & 21 pls. Monné, M.A. (1995) Catalogue of the Cerambycidae (Coleoptera) of the western hemisphere. Part XXII. Subfamily Prioninae. Sociedade Brasileira de Entomologia, São Paulo, XXI, 1–115. Monné, M.A. (2006) Catalogue of the Cerambycidae (Coleoptera) of the Neotropical Region. Part III. Subfamilies Parandrinae, Prioninae, Anoplodermatinae, Aseminae, Spondylidinae, Lepturinae, Oxypeltinae, and addenda to the Cerambycinae and Lamiinae. Zootaxa, 1212, 1–244. Monné, M.A. (2002) Catalogue of the Neotropical Cerambycidae (Coleoptera) with known host plant. Part V: Subfamilies Prioninae, Parandrinae, Oxypeltinae, Anoplodermatinae, Aseminae and Lepturinae. Publicações Avulsas do Museu Nacional, 96, 1–72. Monné, M.A. (2004) Catalogue of the Neotropical Cerambycidae (Coleoptera) with known host plant – part VI: host plant index. Publicações Avulsas do Museu Nacional, 100, 1– 95. Monné, M.A. & Giesbert, E.F. (1994) Checklist of the Cerambycidae and Disteniidae (Coleoptera) of the Western Hemisphere. Wolfsgarden, Burbank, 410 pp. Monné, M.A. & Hovore, F.T. (2005) Checklist of the Cerambycidae, or longhorned beetles (Coleoptera) of the Western Hemisphere. Bio Quip Publications, Rancho Dominguez, 393 pp. Monné, M.A. & Hovore, F.T. (2006) Checklist of the Cerambycidae, or longhorned wood-boring beetles, of the Western Hemisphere. Bio Quip Publications, Rancho Dominguez, 394 pp. Montrouzier, X. (1861) Essai sur la faune entomologique de la Nouvelle-Calédonie (Balade) et des îles des Pins, Art, Lifu, etc. Annales de la Société Entomologique de France, 4(1), 265–306. Neumann, V., Gedeon, K. & Adlbauer, K. (2004) Bockkäferfauna (Coleoptera, Cerambycidae) von Sokotra (Jemen). Mitteilungen der Deutschen Gesellschaft für allgemeine und angewandte Entomologie, 14, 137–140. Noguera, F.A. & Chemsak, J.A. (1996) Cerambycidae (Coleoptera). In: Biodiversidad, Taxonomía y Biogeografía de Artrópodos de México: Hacia una Síntesis de su conocimiento. Universidad Nacional Autonoma de México, 1996, 381–409. Noguera, F.A., Zaragoza-Caballero, S., Chemsak, J.A., Rodríguez-Palafox, A., Ramírez, E., González-Soriano, E. & Ayala, R. (2002) Diversity of the Family Cerambycidae (Coleoptera) of the Tropical Dry Forest of Mexico, I. Sierra

42


SANTOS-SILVA ET AL.

de Huautla, Morelos. Annals of the Entomological Society of America, 95(5), 617–627, 5 figs. Palisot de Beauvois, A.M.F.J. (1805) Insectes recueillis en Afrique et en Amérique, dans les royaumes d'Oware et de Bénin, à Sainte-Domingue et dans les États-Unis, pendant les années 1786–1797. Imprimerie de Fain & Cie, Paris, 1 & 2 [1805], i–xvi + 276, 90 pls. Peck, S.B. (2006) The beetle fauna of Dominica, Lesser Antilles (Insecta: Coleoptera): Diversity and distribution. Insecta Mundi, 20(3–4), 165–209. Peck, S.B. (2009) The beetles of Barbados, West Indies (Insecta: Coleoptera): diversity, distribution and faunal structure. Insecta Mundi, 73, 1–51. Quentin, R.M. & Villiers, A. (1975) Faune de Madagascar (Pars 40). Orstom, Paris, 251 pp. Ramos, J.A. (1946) The insects of Mona Island (West Indies). The Journal of Agriculture of the University of Puerto Rico, 30(1), 1–74, 2 pls. Ramos-Elorduy, J. (2006) Threatened edible insects in Hidalgo, Mexico and some measures to preserve them. Journal of Ethnobiology and Ethnomedicine, 2(51), 10 p. Russo, G. (1930) Contributo alla conoscenza dei Cerambycidae (Coleoptera) della Rep. Dominicana (Antille) con descrizione di una nuova specie. Bollettino del Laboratorio di Zoologia Generale e Agraria del R. Istituto Superiore Agrario di Portici, 24(14), 140–147, 2 figs. Santos-Silva, A. & Martins, U.R. (2005) Novos táxons e notas taxonômicas em Prioninae (Coleoptera, Cerambycidae). Iheringia, Ser. Zool., 95(4), 393–403. Santos-Silva, A. & Galileo, M.H.M (2010) A new genus of Macrotomini (Coleoptera, Cerambycidae, Prioninae). Zookeys, 35, 65–75. Schönherr, C.J. (1817) Synonymia insectorum, oder : Versuch einer Synonymie Aller bisher bekannten Insecten ; nach Fabricii Systema Eleutheratorum &:c. Geordnet. Lewerentzischen Buchdrükerey, Skara, 1(3), xi + 1–506. Skiles, D.D. (1978) Taxonomy and description of two prionine Cerambycidae from southern Arizona: a new species of Stenodontes and new status for Neomallodon arizonicus (Coleoptera). Proceedings of the Entomological Society of Washington, 80(3), 407–423. Smyth, E.G. (1934) The gregarious habit of beetles. Journal of the Kansas Entomological Society, 7(4), 102–119. Tavakilian, G.L. & Chevillotte, H. (1999) Cerambycidae. Available from: https://www.orleans.ird.fr/titan/ (01.07.2009). Thomson, J. (1860–1861) Essai d’une classification de la famille des ce´rambycides et mate´riaux pour servir a` une monographie de cette famille. Paris, 396 pp. + 3 pls. [1860, pp. xvi + 128; 1861, pp. 129–396]. Thomson, J. (1864–65) Systema cerambycidarum ou exposé de tous les genres compris dans la famille des cérambycides et familles limitrophes. Mémoires de la Société Royale des Sciences de Liège, 19, 1– 578 [1864, pp. 1–352; 1865, pp. 353–578]. Thomson, J. (1867) IV. Révision du groupe des Mallodonites (insectes Coléoptères, Prionites, Cérambycides). Physis Recueil d'Histoire Naturelle, (2)1, 85–106. Thomson, J. (1878) Typi cerambycidarum musei Thomsoniani. E. Deyrolle, Paris, 21p. Turnbow, R.H. Jr. & Franklin, R.T. (1980) An annotated checklist of the Cerambycidae of Georgia (Coleoptera). Journal of the Georgia Entomological Society, 15(3), 337–349. Turnbow, R.H., Cave, R.D. & Thomas, M.C. (2003) A list of the Cerambycidae of Honduras, with additions of previously unrecorded species. Ceiba, 44(1), 1–43. Villiers, A. (1980) Coléoptères Cerambycidae des Antilles Françaises I. Parandrinae, Prioninae, Lepturinae. Annales de la Société Entomologique de France (n.s.), 16 (1), 133–157. Vitali, F. (2008) Taxonomic and faunistic notes about the genus Olethrius Thomson, 1860 (Coleoptera Cerambycidae). Entomapeiron (N.S.), 2(1), 1–32. Vitali, F. & Rezbanyai-Reser, L. (2003) Beiträge zur Insektenfauna von Jamaika, Westindien (Karabik) 5. Bockkäfer – I (Coleoptera, Cerambycidae). Les Cahiers Magellanes, 26, 1–16. Wappes, J.E., Morris III, R.F., Nearns, E.H. & Thomas, M.C. (2006) Preliminary checklist of Bolivian Cerambycidae (Coleoptera). Insecta Mundi, 20(1–2), 1–45. Wappes, J.E., Arias, J.L. & Nearns, E.H. (2009) List of Coleoptera Holotypes in Museo de Historia Natural, Noel Kempff Mercado, Universidad Autonoma “Gabriel Rene Moreno,” Santa Cruz de la Sierra, Bolivia. Insecta Mundi, 0081, 1–8. Waterhouse, C.O. (1874) December 7, 1874. Synonymical Notes on Longicorn Coleoptera. The Transactions of the Entomological Society of London, 1874, xxvii–xxix. Waterhouse, C.O. (1881) On the Coleopterous insects collected by Prof. I. Bailey Balfour in the island of Socotra. Proceedings of the Zoological Society of London, 1881, 469–478. White, A. (1853) Catalogue of the coleopterous insects in the collection of the British Museum, pt. VII. Longicornia I. British Museum, London, 7, 1–174. Wolcott, G.N. (1924) “Insectae Portoricensis”. A preliminary annotated checklist of the insects of Porto Rico, with descriptions of some new species. Journal of the Department of Agriculture of Puerto Rico, 7(1), 5–313. Wolcott, G.N. (1936) “Insectae Borinquensis”. A revised annotated checklist of the insects of Porto Rico, with



43

descriptions of some new species. The Journal of Agriculture of the University of Puerto Rico, 20, 1–627. Wolcott, G.N. (1941) A supplement to “Insectae Borinquensis”. The Journal of Agriculture of the University of Puerto Rico, 25(2), 33–158. Wolcott, G.N. (1951) The insects of Puerto Rico. Coleoptera. The Journal of Agriculture of the University of Puerto Rico, 32(2)[1948], 225–416, 56 figs. Zayas, F. (1957) Revisión de los longicórnios Priónidos de Cuba (Coleópteros, Cerambycidae). Memorias de la Sociedad Cubana de Historia Natural “Felipe Poey”, 23, 149–181, 7 pls. Zayas, F. (1975) Revisión de la família Cerambycidae (Coleoptera, Phytophagoidea). Academia de Ciencias de Cuba, Instituto de Zoología, La Habana, 1–433, 140 figs (38 pls).

44


SANTOS-SILVA ET AL.