Associated with Senna Neglecta (Vogel) HS Irwin and Barneby

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Journal of Natural History Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/tnah20 Bruchines (Coleoptera: Chrysomelidae) associated with Senna neglecta (Vogel) H.S. Irwin and Barneby (Fabaceae: Caesalpinioideae): a new host plant for the subfamily Jéssica Herzog Viana a & Cibele Stramare Ribeiro-Costa a a Laboratório de Sistemática e Bioecologia de Coleoptera (Insecta), Departamento de Zoologia , Universidade Federal do Paraná , Curitiba , Brazil Published online: 06 Aug 2013.

To cite this article: Journal of Natural History (2013): Bruchines (Coleoptera: Chrysomelidae) associated with Senna neglecta (Vogel) H.S. Irwin and Barneby (Fabaceae: Caesalpinioideae): a new host plant for the subfamily, Journal of Natural History To link to this article: http://dx.doi.org/10.1080/00222933.2013.791882

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Bruchines (Coleoptera: Chrysomelidae) associated with Senna neglecta (Vogel) H.S. Irwin and Barneby (Fabaceae: Caesalpinioideae): a new host plant for the subfamily Jéssica Herzog Viana* and Cibele Stramare Ribeiro-Costa

Laboratório de Sistemática e Bioecologia de Coleoptera (Insecta), Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Brazil

(Received 14 September 2011; ﬁnal version received 29 March 2013)

The structures of the exoskeleton and genitalia of bruchines preying on Senna neglecta (Vogel) H.S. Irwin and Barneby seeds were studied. Three new species are described and S. neglecta is recorded as a new host plant for the subfamily. The six bruchine species associated with S. neglecta are Amblycerus hoffmanseggi (Gyllenhal 1833) (Amblycerini), Acanthoscelides multimaculatus sp. nov., Sennius bondari (Pic 1929), Sennius nappi Ribeiro-Costa and Reynaud 1998, Sennius margarete sp.nov.andSennius ricardo sp. nov. (Bruchini). Sennius bondari is the most polyphagous species, while Sennius margarete sp.nov.andSennius ricardo sp. nov. are currently only known from S. neglecta. Descriptions, redescriptions or diagnoses are presented, including revised characters based on electron microscope images of the denticles at the base of the elytra striae, and the lateroventral carina of the hind tibia. An identiﬁcation key, illustrations, a list of host plants and geographical distribution records are also included. http://www.zoobank.org/urn:lsid:zoobank.org:pub:4293CF55-AA92-4FD5-8343- E51BB9EE5ABD Keywords: Amblycerus; Acanthoscelides; Sennius; seed-beetles; Cassieae

Introduction Senna Mill. is a large genus with about 350 species distributed mainly within the trop- ical Americas (Marazzi et al. 2006). A total of nine genera of bruchines (Coleoptera: Chrysomelidae) have already been recorded as predators on the seeds of this legume (e.g. Kingsolver 1971; Arora 1977; Johnson 1989; Delobel et al. 1995; Casari and Texeira 1997; Ribeiro-Costa 1998; Romero and Johnson 2000, 2004). Senna neglecta (Vogel) H.S. Irwin & Barneby occurs in northeastern, southeast- ern, and southern Brazil (Irwin & Barneby 1982) in ﬁelds, secondary vegetation, and forest edges and interiors (Rodrigues et al. 2005). Studies of this species have focused

Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 almost exclusively on their use in recuperating degraded areas, landscaping and drug production (Leal et al. 2004; Souza et al. 2004; Leal and Biondi 2006). There are no previous records in the literature of seed predation by bruchines and the knowledge of the diversity of seed beetles in this plant is important because these insects can limit successful seed production and therefore population propagation. Earlier unpublished ecological studies conducted by the ﬁrst author with S. neglecta var. oligophylla in the Serra dos Órgãos National Park in the state of Rio

*Corresponding author. Email: [email protected]

de Janeiro, Brazil, and by the second author with S. neglecta var. neglecta in Curitiba, state of Paraná, recorded Amblycerus hoffmanseggi (Gyllenhal 1833) and Sennius nappi Ribeiro-Costa & Reynaud 1998, as predators of their seeds, although it was not possible at that time to identify many specimens to the specific level. The present study reviewed the bruchines associated with S. neglecta and provides specific names and an identification key to facilitate their recognition.

Material and methods The material studied came mainly from collections made on seeds of S. neglecta var. oligophylla at the Serra dos Órgãos National Park, Rio de Janeiro State, Brazil (22◦4518 S, 43◦0020 W), and in Curitiba, Paraná State (25◦4503 S, 49◦2426 W) on S. neglecta var. neglecta seeds. Other specimens (or their photographs) were acquired on loan; the new species have been deposited in different national and international museums, these are listed below followed by the acronyms and the name of curator: BMNH, British Museum of Natural History, London, UK (Photos: S.P. Rosa); CEAM, Centro de Entomologia y Acarologia, Montecillo, Mexico (J.R. Nápoles); CNCI, Canadian National Collection of Insects, Ottawa, Canada (A.E. Davies); DZUP, Coleção de Entomologia Pe. J.S. Moure, Curitiba, Paraná, Brazil (C.S. Ribeiro-Costa); FIOC, Fundação Instituto Oswaldo Cruz, Rio de Janeiro, Rio de Janeiro, Brazil (S. de Oliveira); FSCA, Florida State Collection of Arthropods, Gainesville, FL, USA (M.C. Thomas); MCNZ, Museu de Ciências Naturais da Fundação Zoobotânica do Rio Grande do Sul, Porto Alegre, Rio Grande do Sul, Brazil (M.H. Galileo); MNRJ, Museu Nacional/Universidade Federal do Rio de Janeiro, Rio de Janeiro, Rio de Janeiro, Brazil (M.A. Monné); MZSP, Museu de Zoologia, Universidade de São Paulo, São Paulo, Brazil (S. Casari); TAMU, Texas A&M University, College Station, TX, USA (E. Riley); USNM, National Museum of Natural History, Washington, D. C., USA (D.G. Furth). The date format of non-type specimens was standardized regardless of their label information to: day.month (in Roman numerals).year. Most characters were observed from dry and pinned insects. When necessary, specimens were boiled in water for 5 minutes and dissected under a LEICA MTU253 stereoscopic microscope. Drawings were made using the same equipment. To study the male genitalia, the pygidium was raised and the internal mass of abdominal muscles and organs, including the genitalia, was removed and boiled in 10% potassium hydroxide for a few minutes. This material was then washed in distilled water, placed on a slide, and observed under a Carl Zeiss–Standard 20 microscope; images were captured using a Sony Cyber-Shot DSC–W300 digital camera coupled

Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 to the microscope; drawings of the internal sac sclerites were made with the same equipment. Coloured images of the external morphology were captured with a LEICA DFC 500 digital camera (IMAGE MANAGER) attached to a LEICA MZ16 stereomicro- scope and subsequently processed using Auto-Montage Pro (Syncroscopy, Cambridge, United Kingdom) image processing software of the “TAXon line – Rede Paranaense de Coleções Biológicas” at the Departamento de Zoologia, UFPR. Images were made of non-metalized specimens at low vacuum levels using a Jeol JSM-6360LV scanning electron microscope at the Centro de Microscopia Eletrônica (CME) of UFPR. Journal of Natural History 3

The terminology adopted follows Johnson and Kingsolver (1973), with the description of the internal sac of male genitalia in the everted position, and Silva et al. (2003). The host plants and the geographic distributions of each species were based on label data, the published literature and the Brucol Database (Romero & Johnson 2002), summarized in Table 1. All plant names were checked using the International Legume Database and Information Service (ILDIS) and grouped according to their current classiﬁcation into family (bold), subfamily (underlined) and species, in alphabetical order. The following abbreviations were used for countries: ARG, Argentina; BIZ, Belize; BOL, Bolivia; BRA, Brazil; CHI, Chile; COL, Colombia; CRC, Costa Rica; DOM, Dominican Republic; ECU, Ecuador; ESA, El Salvador; GRN, Grenada; GUA,

Table 1. List of bruchines associated with Senna neglecta with their host plants and country distribution.

Bruchinae associated Host plants Geographic distribuition with Senna neglecta

Acanthoscelides Senna neglecta var. oligophylla, Senna BRA multimaculatus neglecta var. neglecta and Senna sp. n occidentalis (2 spp.) Amblycerus Gleditshia triacanthos, Senna alata, BRA (NDS), HAI, DOM, hoffmanseggi S. bicapularis, S. occidentalis, ARG, BOL, PAR S. corymbosa, S. aphylla, S. bauhinioides, S. hirsuta, S. neglecta var. oligophylla (NHP), S. neglecta var. neglecta (NHP) and Macroptilium atropurpureum (10 spp.) Sennius bondari Memora peregrina, Senna BRA (NDS), VEN, COL, pistacilfolia, S. pendula, BOL S. occidentalis, S. bicapsularis, S. macranthera, S. multijuga, S. surattensis, S. splendida, S. alata, S. neglecta var. oligophylla (NHP), Senna neglecta var. neglecta (NHP) and S. appendiculata (NHP) (12 spp.) Sennius margarete Senna neglecta var. oligophylla and BRA sp. nov. Senna neglecta var. neglecta (1 sp.)

Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 Sennius nappi Senna multijuga, S. neglecta var. BRA (NDS) oligophylla (NHP), S. neglecta var. neglecta (NHP), S. rugosa (NHP), S. occidentalis (NHP) and S. obtusifolia (NHP) (5 spp.) Sennius ricardo sp. Senna neglecta var. oligophylla (1 sp.) BRA nov. Notes: NHP, new record of host plant; NDC, new geographical distribution to country; NDS, new geographical distribution to states. 4 J. Herzog Viana and C.S. Ribeiro-Costa

Guatemala; HAI, Haiti; HON, Honduras; JAM, Jamaica; MEX, Mexico; NCA, Nicaragua; PAN, Panama; PAR, Paraguay; PUR, Puerto Rico; TRI, Trinidad and Tobago; USA, United States of America; VEN, Venezuela.

Results and discussion We identiﬁed six bruchine species associated with Senna neglecta: Amblycerus hoffmanseggi (Gyllenhal 1833) (Amblycerini), Acanthoscelides multimaculatus sp. nov. (Bruchini), Sennius bondari (Pic 1929) (Bruchini), Sennius nappi Ribeiro-Costa and Reynaud 1998 (Bruchini), Sennius margarete sp. nov. (Bruchini) and Sennius ricardo sp. nov. (Bruchini).

Key to the bruchine species associated with Senna neglecta 1. Shallow ocular sinus (Figure 2C). Subtriangular scutellum (Figure 2A). Pygidium slightly covered by elytra (Figure 2B, D). Hind leg with femur without teeth on ventral margin and tibia with a pair of spurs (Figure 2E, F) (Genus Amblycerus)...... Amblycerus hoffmanseggi Distribution: ARG, BOL, BRA, DOM, HAI, PAR. Host plants: Gleditsia triacanthos, Senna alata, S. aphylla, S. bauhinioides, S. bicapularis, S. corymbosa, S. hirsuta, S. neglecta var. oligophylla, S. neglecta var. neglecta, S. occidentalis, Platypodium elegans, Macroptilium atropurpureum. Deep ocular sinus (Figure 1C). Subquadrate scutellum (Figure 1A). Pygidium completely exposed (Figure 1B, D). Hind leg with at least one tooth on ventral margin of femur and tibia without spurs (Figure 1F, G)...... 2 2. Hind femur with one long tooth followed by two other smaller on ventral margin, near the apex (Figure 1F). Male genitalia without hinge sclerites (Genus Acanthoscelides)...... Acanthoscelides multimaculatus sp. nov. Distribution: BRA. Host plants: Senna neglecta var. oligophylla, S. neglecta var. neglecta, S. occidentalis. Hind femur with only one smooth or serrated tooth on ventral margin, near the apex (Figure 3F). Male genitalia with hinge sclerites (Genus Sennius)...3 3. Elytra usually with pubescence evenly distributed (Figure 4A, E, I); dense pubescence, when present, only at base of elytra (Figure 4I). Male genitalia with hinge sclerites almost straight (Figure 4P) . . . Sennius margarete sp. nov. Distribution: BRA. Host plants: Senna neglecta var. oligophylla, S. neglecta var. neglecta. Elytra never with pubescence evenly distributed (Figure 3A), with dense white Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 bands of vertical and horizontal pubescence, usually at base. Male genitalia with hinge sclerites curved (Figure 3H)...... 4 4. Elytron with wide semicircular macula of white pubescence over most of the base and the apex of the second striae interval at its smaller width, reaching its greatest width at the middle of the ﬁfth striae interval (Figure 3A). Male genitalia with hinge sclerites broad (Figure 3H) ...... S. bondari Distribution: COL, VEN, BOL, BRA. Host plant: Memora peregrina, Senna pistacilfolia, S. pendula, S. occidentalis, S. bicapsularis, S. macranthera, Journal of Natural History 5

S. multijuga, S. surattensis, S. splendida, S. alata, Senna neglecta var. oligophylla, S. neglecta var. neglecta, S. australis. Elytron with transverse submedian band of white pubescence covering at least the second to ninth striae intervals, usually over a red-orange to dark brown macula (Figure 5A). Male genitalia with hinge sclerites narrow (Figure 5H) ...... 5 5. Pronotum with heavy, dense, white pubescence in short oblique band at distal corners (Figure 5A). Hind femur with a slightly prominent and microserrate tooth (Figure 5F). Male genitalia with long hinge sclerites (Figure 5H) ...... S. nappi Distribution: BRA. Host plants: Senna multijuga, Senna neglecta var. oligophylla, S. neglecta var. neglecta, S. rugosa, S. occidentalis,S. obtusifolia. Pronotum with moderately dense white pubescence at distal corners (Figure 6A). Hind femur with a very prominent and strongly serrate tooth (Figure 6E). Male genitalia with short hinge sclerites (Figure 6G) ...... Sennius ricardo sp. nov. Distribution: BRA. Host plant: Senna neglecta var. oligophylla.

Acanthoscelides multimaculatus sp. nov. (Figure 1A–I)

Description Dimension. Length (Pronotum-elytra) 2.32–2.96 mm. Width 1.48–1.92 mm.

Integument colour. Head, thorax and abdomen dark brown to black (Figure 1B). Antenna dark brown or with the ﬁrst four segments light brown (Figure 1B, C). Maxillary and labial palps black (Figure 1C). Elytron dark brown with light brown oblique macula on the basal half, from base of the fourth to seventh striae intervals until second to ﬁfth striae intervals in the median region; additional macula at part of the posterior margin in the apical region (Figure 1A). Pygidium light brown to black (Figure 1D). Legs light brown, except middle coxa and basal two-thirds of hind femur dark brown, and tarsus red-orange (Figure 1B).

Pubescence. Head with dense tuft of white hairs posterior to postocular lobe (Figure 1C). Ocular sinus, postocular lobe, and mandibles with moderately dense white pubescence (Figure 1C). Labrum aligned with golden pubescence near the base and apex (Figure 1C). Pronotum with sparse pubescence and small tufts of intermixed

Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 white with golden pubescence at base and lateral thirds of apex and at antescutellar region (Figure 1A). Scutellum with sparse white pubescence (Figure 1A). Elytron with white pubescence strongly dense only in a short band at submedian region of third striae interval, remaining portions with white and golden pubescence moderately dense, distributed as follows: macula elongate and oblique at the anterior half that covers part of the striae intervals fourth to seventh; on small, short and slightly oblique band at the submedian region that covers part of the seventh to ninth striae intervals; on broad and irregular macula in the posterior half that covers the second to ninth striae intervals; remaining elytron with golden pubescence (Figure 1A). Pygidium with 6 J. Herzog Viana and C.S. Ribeiro-Costa

Figure 1. Acanthoscelides multimaculatus sp. nov. (A) Dorsal view; (B) lateral view; (C) head, frontal view; (D) pygidium; (E) base of elytra striae; (F) part of hind leg, internal view; (G) hind tibia, external view; (H, I) male genitalia: (H) median lobe with sclerites in armature of internal sac magniﬁed; (I) lateral lobes. DMC, dorsomesal carina; LC, lateral carina; LVC, lateroventral carina; VC, ventral carina; M, mucro; BR, basal region; MR, median region; AR, apical region; VV, ventral valve.

white and golden sparse pubescence; dense white pubescence at middle, and moderately dense in basal region, lateroanterior areas and median line (Figure 1D). Ventral surface with white pubescence denser at the base of mesepimeron, metepisternum, and tufts on the lateral areas of the abdominal ventrites (Figure 1B). Head evenly punctuate, except in the midline of frons with only fine punctuation, apex of clypeus and labrum smooth; frons convex, frontal carina and transverse sulcus absent; ocular sinus deep, more than two-thirds eye length; postocular lobe narrow (Figure 1C). First and third antenna segments filiform, second and fourth moniliform, fifth to tenth wider than long, and eleventh globular with pointed apex.

Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 Pronotum moderately convex; disc slightly sulcate at basal lobe (Figure 1A). Scutellum subquadrate, slightly wider than long (Figure 1A). Striae third and fourth at base with denticles (Figure 1E). Hind femur with two small teeth (0.02–0.04 mm), and one more prominent (0.06–0.10 mm) with wide base on the ventral margin, near the apex (Figure 1F). Hind tibia with lateroventral carina (Figure 1G – LVC) extending more than half of its length; lateral (Figure 1G – LC), dorsomesal (Figure 1F – DMC) and ventral (Figure 1G – VC) carinae long, ending near coronal denticles; mucro (Figure 1F – M) 2.5 times longer than the opposite coronal denticle. Last abdominal ventrite of male emarginate at middle, rounded in female. Pygidium not concealed by Journal of Natural History 7

elytra, lateral margins slightly curved, with apex rounded in male, more acuminate in female (Figures 1B, D).

Male genitalia. Median lobe about 4.3 times longer than its width in median region, expanded at the apex; ventral valve rounded, with convex lateral margins. Internal sac not apically lobed, basal region without spicules near the base of ventral valve; median region with wide sclerite in the form of an inverted “V”, with microserrate margin at apex and ﬂanked by two pairs of sclerites, the anterior being long and thin and pointed at apex, nearly as long as the “V” sclerite, posterior sclerites short, with wide bases similar to teeth (Figure 1H). Tegmen with lateral lobes separated by deep emargination, about 0.63 times the length of lateral lobes (Figure 1I).

Type material Holotype, allotype and 10 paratypes deposited in DZUP with labels: “Brasil, RJ – Teresópolis | P.N. Serra dos Órgãos | 22/VIII/2006 | J.H. Viana col.” “Em | Senna neglecta | var. oligophylla”. Five paratypes with the same label as the holotype deposited at each of the following museums: MZSP, MNRJ, TAMU, USNM, FSCA, CEAM and CNCI.

Additional specimens BRAZIL: Rio de Janeiro: Teresópolis: P.N. Serra dos Órgãos, VII/2006, J.H. Viana col., in Senna neglecta var. oligophylla, 48 specimens (DZUP); 22/VIII/2006, same collector and host plant, 19 specimens (DZUP); 05/IX/2006, same collector and host plant, 97 specimens (DZUP); 21/IX/2006, same collector and host plant, 15 specimens (DZUP); 22/VIII/2007, same collector and host plant, 1 specimen (DZUP); 06/IX/2007, same collector and host plant, 8 specimens (DZUP); 12/IX/2007, same collector and host plant, 7 specimens (DZUP); 14/IX/2007, same collector and host plant, 3 specimens (DZUP); 21/IX/2007, same collector and host plant, 6 specimens (DZUP); 25/X/2007, same collector and host plant, 1 specimen (DZUP). Mato Grosso do Sul: Campo Grande: 16/VIII/[19]87, F. Giacomel col., no host plant, 21 specimens (DZUP). Paraná: Curitiba: 9/VII/1997, C.S. Ribeiro-Costa col., Senna neglecta var. neglecta, 3 specimens (DZUP); 28/VII/1997, same collector and no host plant, 1 specimen (DZUP); 20/IX/2001, E. Caron col., no host plant, 2 specimens (DZUP); 28/IX/2001, same collector and no host plant, 2 specimens (DZUP); 02/X/2001, same collector and no host plant, 3 specimens (DZUP); 05/X/2001, same collector and no host plant, 1 specimen (DZUP); 10/X/2001, same collector and no host plant, 2 specimens (DZUP); 11/X/2001, same collector and no host plant, 2 spec-

Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 imens (DZUP); 12/X/2001, same collector and no host plant, 2 specimens (DZUP); 15/X/2001, same collector and no host plant, 5 specimens (DZUP); 26/X/2001, same collector and no host plant, 1 specimen (DZUP); 04/XI/2001, same collector and no host plant, 1 specimen (DZUP). Londrina: 10/VI/2002, A. Menezes Jr. col., Senna occidentalis, 2 specimens (DZUP). No locality: no date or collector, 1 specimen (DZUP).

Distribution Brazil (Rio de Janeiro, Mato Grosso do Sul, and Paraná). 8 J. Herzog Viana and C.S. Ribeiro-Costa

Host plants Fabaceae: Caesalpinioideae: Senna neglecta var. oligophylla, S. neglecta var. neglecta, S. occidentalis.

Discussion This species differs from others found on Senna neglecta seeds mainly by the light pubescence on dark integument forming a variegated pattern on almost the entire elytra (Figure 1A), by the presence of a prominent tooth and two smaller teeth at the inner margin of hind femur (Figure 1F), and absence of hinge sclerites in the median lobe of the male genitalia, which are diagnostic for Sennius species (Figure 3H – HS). Acanthoscelides multimaculatus sp. nov. belongs to the quadridentatus group of Acanthoscelides established by Kingsolver (1980) along with 15 other species by sharing with them similar pattern of the male genitalia. The species are A. devriesi Kingsolver 1980, A. difficilis (Sharp, 1885), A. equivocada Johnson 1990, A. laicus (Fahraeus 1839), A. machala Johnson 1990, A. mimosicola Johnson 1983, A. pectoralis (Horn 1873), A. pigricola Kingsolver 1980, A. puniceus Johnson 1983, A. quadridentatus (Schaeffer 1907), A. ramirezi Johnson 1990, A. tridenticulatus Bottimer 1969, A. triumfettae Kingsolver 1980, A. zebratus Kingsolver 1980 and A. zulia Johnson 1990. Of these species, 11 feed exclusively on the genus Mimosa Linnaeus (Fabaceae; Mimosoideae), one (A. pectoralis) is associated with Desmanthus Willdenow (Fabaceae: Mimosoideae) and one other (A. triumfettae) with Triumfetta Linnaeus (Malvaceae), however A. laicus, A. machala and A. multimaculatus sp. nov. feed on Senna (Fabaceae: Caesalpinioideae). Acanthoscelides multimaculatus sp. nov. seems more similar to A. machala by the five sclerites of the internal sac of the male genitalia The differences between them rely mainly on the margin of sclerite shaped like an inverted “V” in the male genitalia, which is microserrate in A. multimaculatus sp. nov. and strongly serrate in A. machala (Johnson 1990, figs 299–303, p. 572).

Etymology The speciﬁc name “multimaculatus” is related to the variegated pubescence pattern on dorsum.

Amblycerus hoffmanseggi (Gyllenhal, 1833) (Figure 2A–H)

Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 Spermophagus hoffmanseggi Gyllenhal 1833, p. 105 (original description, distribution); Dejean 1837, p. 255 (catalogue, distribution); Schöenherr 1839, p. 134 (citation, distribution); Jekel 1855, p. 30 (citation); Gemminger and Harold 1873, p. 3218 (catalogue, distribution); Sharp 1885, p. 496 (citation); Pic 1913, p. 59 (catalogue, distribution); Pierce 1915, p. 9 (citation); Leng 1920, p. 306 (catalogue, distribution); Blair 1928, p. 679 (citation); Costa Lima 1928, p. 183 (host, distribution); Bondar 1931, p. 59 (citation, distribution); 1936, p. 34 (citation, distribution); Bosq 1943, p. 47 (distribution, host); Zacher 1952, p. 468 (host); Guérin 1953, p. 179 (citation, distribution, redescription); Kingsolver and Silva 1991, p. 414 (citation). Journal of Natural History 9

Figure 2. Amblycerus hoffmanseggi. (A) Dorsal view; (B) lateral view; (C) head, frontal view; (D) pygidium; (E) part of hind leg, internal view; (F) hind tibia, external view; (G, H) male genitalia: (G) median lobe with sclerites in armature of internal sac magniﬁed; (H) lateral lobes.

Amblycerus hoffmanseggi: Bridwell 1944; Blackwelder 1946, p. 762 (catalogue, distribution); Costa Lima 1955, p. 249 (host); Silva et al. 1968, p. 376 (distribution, host); Kingsolver 1970, p. 485 (redescription, distribution, male genitalia figure, key); Terán and L’Argentier 1981, pp. 61–70 (immature figure, ecology, host); Johnson and Kingsolver 1982, p. 410 (catalogue, distribution); Borowiec 1987, p. 187 (male genitalia figure); Udayagiri and Wadhi 1989, p. 10 (catalogue, distribution, type locality); Ribeiro-Costa and Marinoni 1992, pp. 149–175 (phenetic analisys); Ribeiro-Costa 1992, p. 153 (redescription, distribution, key, biology, host, male genitalia figure); Macêdo et al. 1992, pp. 331, 333 (biology, host); Johnson et al. 2001, pp. 37, 44, 45 (citation); Kingsolver 2004, p. 35 (citation). Bruchus robiniae Olivier 1795, p. 7 (nec Fabricius, 1781, p. 75) (redescription); Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 Gyllenhal 1833, p. 105 (synonym).

Diagnosis Dimensions. Length (Pronotum-elytra) 4.04–5.35 mm. Width 2.36–3.12 mm.

Integument colour. Pronotum and elytra usually dark brown to black (Figure 2A). Antennae, edges and ventral region of pronotum dark brown (Figure 2A, B). Elytron 10 J. Herzog Viana and C.S. Ribeiro-Costa

with black longitudinal band on the external margin, rarely dark brown, from the tenth and eleventh or ninth to eleventh, or eighth to eleventh striae intervals (Figure 2A). Pygidium evenly dark brown (Figure 2D) or sometimes orange-red in lateroanterior areas. Abdominal ventrites usually with lateral rounded areas orange-red, rarely with the same colour (Figure 2B).

Pubescence. General pattern – Dense tuft of white pubescence posterior to the postocular lobe absent. Pronotum with dense yellow pubescence distributed in tufts, without forming distinct longitudinal bands or, sometimes, with dense pubescence forming a median band generally not reaching the anterior margin; white pubescence finely bor- dering edges; black pubescence scattered or in rare tufts (Figure 2A). Scutellum with dense white pubescence (Figure 2A). Elytron with intermixed light yellow and white pubescence with black tufts, the white and light yellow pubescence are distributed as follows: white on the inner margin, light yellow from the first to seventh striae intervals, white on eighth to eleventh, or light yellow on first to eighth and white on ninth to eleventh striae intervals, both intermixed with black tufts (Figure 2A). Pygidium with white pubescence denser at basal and lateral areas (Figure 2D). Pattern B – Pronotum with dense white pubescence laterally in tufts and, sometimes, on the midline forming a prescutellar band; rest with black scattered pubescence. Elytra with intermixed white pubescence and black tufts. Ocular sinus shallow, not more than one-third of eye length (Figure 2C). Pronotum slightly convex, disc not sulcate in basal lobe (Figure 2A). Scutellum subtriangular (Figure 2A). Hind femur without teeth (Figure 2E). Hind tibia without carina and mucro (Figure 2F), a pair of spurs present, with internal spur about half the length of the external (Figure 2E, F). Last abdominal ventrite of male and female not emarginate at median region. Pygidium slightly concealed by elytra, lateral margins slightly straight with rounded apex (Figures 2B, D).

Male genitalia. Median lobe about ﬁve times longer than its median width, slightly expanded at the apex. Ventral valve (Figure 2G – VV) elongate and triangular, lateral margins sinuous near the opening of the internal sac. Basal region of internal sac (Figure 2G – BR) without spicules near base of ventral valve, with an anterior pair of small sclerites similar to teeth, posterior elongated, about six times its width, with apex rounded in ventral view; median region (Figure 2G – MR) with pair of lateral falciform sclerites with apexes varying in curvature, between them a median falciform sclerite about half the length of the lateral sclerites (Figure 2G). Tegmen with lateral lobes separated by shallow emargination, about 0.1 times the length of lateral lobes (Figure 2H). Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 Type material According to Udayagiri and Wadhi (1989), the holotype is deposited at Swedish Museum of Natural History, Stockholm, Sweden (NHRS). Labels: “typus [red]” “Br: Robinia, Rio de Janeiro, V. Winther [handwritten, white]” “282, 69 [rose]”. In 1970, Kingsolver examined the type and redescribed Am. hoffmanseggi.Later, when Ribeiro-Costa (1992) established the hoffmanseggi group of Amblycerus,she redescribed this species again indicating the patterns of coloration, besides illustrating the male genitalia. Journal of Natural History 11

Additional specimens BRAZIL: Rio de Janeiro: Arraial do Cabo: 10/X/1987, L.R.C. Souza col., in fruit of Senna bicapsularis, 1 specimen (DZUP). Itatiaia: 5/X/1924, Zikán col., no host plant, 1 specimen (FIOC). Teresópolis: P.N. Serra dos Órgãos, 2006, J.H. Viana col., in Senna neglecta var. oligophylla, 3 specimens (DZUP); VI/2006, same collector and host plant, 43 specimens (DZUP); 22/VIII/2006, same collector and host plant, 12 specimens (DZUP); 05/IX/2006, same collector and host plant, 8 specimens (DZUP); 2007, same collector and host plant, 1 specimen (DZUP); 30/VII/2007, same collector and host plant, 1 specimen (DZUP); 12/X/2007, same collector and host plant, 1 specimen (DZUP); 14/IX/2007, same collector and host plant, 2 specimens (DZUP); 21/IX/2007, same collector and host plant, 1 specimen (DZUP); 26/IX/2007, same collector and host plant, 1 specimen (DZUP). Paraná: Curitiba: II/1988, F. Giacomel col., Cassia alata, 2 specimens (DZUP); II/1988, same collector, 1 specimen (DZUP); 02/V/1997, C.S. Ribeiro-Costa e J.M. Bobato col., Senna neglecta, 15 specimens (DZUP); VI/1997, C.S. Ribeiro-Costa col., Senna neglecta var. neglecta, 9 specimens (DZUP); VI/1998, same collector and host plant, 28 specimens (DZUP); Jardim Botânico, 20/V/2003, no collector and same host plant, 37 specimens (DZUP); UFPR – Prédio Ed. Física, 06/VII/2008, no collector and same host plant, 7 specimens (DZUP). Prudentópolis:1/XII/1968, Moure & Dressler col., no host plant, 1 specimen (DZUP). Santa Catarina: Mafra: 08/I/[19]93, D.T.R. Santos col., no host plant, 1 specimen (DZUP). Rio Grande do Sul: Viamão: 24/II/1986, M. Hoffmann col., 1 specimen (MCNZ). No locality: 5/X/[19]24 | i. [not legible], Fedegoso, 1 specimen (FIOC).

Distribution United States of America (erroneous record, Kingsolver 1970), Haiti and the Dominican Republic (erroneous record, Kingsolver 1970), Brazil (Bahia, Ceará, Espírito Santo, Pará, Mato Grosso, Goias, Minas Gerais, Rio de Janeiro, São Paulo, Paraná), Bolivia, Paraguay (San Pedro), Argentina (Corrientes, Entre Rios, Jujuy, Misiones, Salta, Tucuman). New Records. Brazil (Santa Catarina, Rio Grande do Sul).

Host plants Fabaceae: Caesalpinioideae: Gleditsia triacanthos, Senna alata, S. bicapularis, S. occidentalis, S. corymbosa, S. aphylla, S. bauhinioides, S. hirsuta; Faboideae: Platypodium elegans (doubtful record, Ribeiro-Costa 1992), Macroptilium atropurpureum.

Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 Malvaceae: Malvoideae: Gossypium sp. (doubtful record, Ribeiro-Costa 1992). New records – Fabaceae: Caesalpinioideae: Senna neglecta var. oligophylla and Senna neglecta var. neglecta.

Discussion This species differs from all of the other bruchines occurring on S. neglecta by its larger size and by having a pair of spurs on the hind tibia. It has three pubescence colour patterns (Ribeiro-Costa 1992), although we found only two of them (the general pattern 12 J. Herzog Viana and C.S. Ribeiro-Costa

and pattern B) associated with S. neglecta. The general pattern was the most frequent in S. neglecta, and is presented and illustrated here; pattern B was the rarest, appearing in only one specimen. Later, Ribeiro-Costa (1992) established the hoffmanseggi group of Amblycerus and redescribed this species again, indicating the patterns of coloration and illustrating the male genitalia.

Sennius bondari (Pic, 1929) (Figure 3A–I)

Bruchus bondari Pic 1929, p. 28. (original description, distribution); Bondar 1936, p. 40 (redescription, host, distribution); Zacher 1952, p. 461 (host). Acanthoscelides bondari: Blackwelder 1946, p. 759 (catalogue, distribution); Costa Lima 1955, p. 245 (host); Silva et al. 1968, p. 372 (host, distribution). Sennius bondari: Johnson and Kingsolver 1973, p. 72 (citation); Johnson 1984, p. 62 (host, distribution); Udayagiri and Wadhi 1989, p. 100 (catalogue, distribution, host); Kingsolver & Silva 1991, p. 412 (citation); Macêdo et al. 1992, pp. 331, 334 (host, distribution); Pimentel 1997 (biology, host); Ribeiro-Costa and Reynaud 1998, p. 245 (catalogue, distribution, host); Ribeiro-Costa 1998, p. 66 (biology); Silva et al. 2003, p. 269 (citation); Linzmeier et al. 2004, p. 351 (biology); Caron et al. 2004, p. 4 (egg description); Koller and Nunes 2006, p. 6 (host). Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013

Figure 3. Sennius bondari. (A) Dorsal view; (B) lateral view; (C) head, frontal view; (D) pygidium; (E) base of elytra striae; (F) part of hind leg, internal view; (G) hind tibia, external view; (H, I) male genitalia: (H) median lobe with hinge sclerites (HE) and others sclerites in internal sac magniﬁed; (I) lateral lobes. BR: basal region; SMR: submedian region; SAR: subapical region; AR: apical region. Journal of Natural History 13

Redescription Dimension. Length (Pronotum-elytra) 1.96–2.60 mm. Width 1.28–1.64 mm.

Integument colour. Black body (Figure 3A), except labrum, maxillary and labial palps dark brown to black (Figure 3C). Antennae orange-red, some specimens with seven apical segments dark brown (Figure 3B). Legs light to dark brown (Figure 3B).

Pubescence. Head with dense tuft of white pubescence posterior to the postocular lobe (Figure 3C). Ocular sinus, postocular lobe and mandibles with white pubescence moderately dense. Labrum with scattered golden pubescence, denser at the apex (Figure 3C). Pronotum with intermixed white and golden pubescence, moderately white pubescence distributed in small tufts in lateral third and at midline of basal lobe (Figure 3A). Scutellum with dense white pubescence (Figure 3A). Elytron with white, golden and black intermixed pubescence, white pubescence densely distributed as follows: in a continuous line at first striae interval, in small tufts at bases of the third and fifth striae intervals; wide semicircular macula almost covering the base and apex of second striae interval at its smaller width, and reaching its greatest width over middle of fifth striae interval; in small, short and transversal band in the submedian region that covers part of the sixth to ninth striae interval, sometimes dense tuft in seventh and/or ninth striae interval (Figure 3A). Pygidium with intermixed sparse white and golden pubescence; macula of dense white pubescence in the median-basal region and moderately dense in the lateroanterior corners (Figure 3D). Ventral surface with white pubescence denser at base of mesepimeron, posterior margin of metepisternum, distal region of hind coxa, and at base of first abdominal ventrite (Figure 3B). Head with coarse punctuation, except median line of frons with fine punctuation; labrum smooth, frons convex, frontal carina and transverse sulcus absent; ocular sinus deep, more than half an eye length (Figure 3C). First and third antenna segments filiform, second and fourth moniliform, fifth to tenth wider than long, and eleventh globular with pointed apex (Figure 3B). Pronotum moderately convex (Figure 3B); disc with basal lobe not sulcate (Figure 3A). Scutellum subquadrate, slightly wider than long (Figure 3A). Fourth striae of elytron with denticle at base (Figure 3E). Hind femur with a prominent tooth (0.01–0.06 mm) on ventral margin slightly microserrate (Figure 3F). Hind tibia with lateroventral carina extending half its length (Figure 3G), lateral, dorsomesal and ventral carinae long, ending near coronal denticles (Figure 3F, G); mucro about 1.8 times longer than the opposite coronal denticle (Figure 3F, G). Last abdominal ventrite of male emarginate at middle, rounded in female. Pygidium not concealed by elytra, lateral margins slightly curved, with rounded apex in males, more acuminate in females (Figure 3B, D). Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013

Male genitalia. Median lobe about 5.2 times longer than the width at its middle, expanded at apex; ventral valve with convex lateral margins and truncate apex; hinge sclerites long and wide, with external margin moderately concave and sinuous (Figure 3H – HS). Internal sac trilobate, basal region without spines (Figure 3H – BR), submedian region (Figure 3H – SMR) with small spicules in two irregular dense clusters; subapical region (Figure 3H – SAR) with lateral clusters of large spicules not aligned at bases of lateral lobes; apexes of lateral lobes with minute spicules; 14 J. Herzog Viana and C.S. Ribeiro-Costa

apical region (Figure 3H – AR) of middle lobe with denticles near the gonopore sclerite (Figure 3H). Tegmen with lateral lobes separated by deep emargination, about 0.7 times the length of lateral lobes (Figure 3I).

Type material It was possible to compare the type deposited in BMNH only by photos. Labels: “Type” “Brazil | Bahia | 1928 | Dr. G. Bondar. | 1233 [right vertical label]” “Pres. by | Imp. Bur. Ent. | Brit. Mus. | 1928-504” “[unreadable]” “Bruchus | bondari | nsp.” Note: In the original description, Pic (1929) did not mention the number of specimens studied, however, the information on labels of the photographed material correspond to those described in that work, therefore this is probably type material.

Additional specimens COLOMBIA: Valle del Cauca: 10 km W Cali, 17/VII/[19]82, C.D. Johnson col., in seeds of Senna pistacilfolia, 4 specimens (USNM) and 11 specimens (TAMU). Palmira: 5 km E.[Leste], 16/VII/[19]82, C.D. Johnson col., in seeds of Senna pendula var. advena, 7 specimens (TAMU); BRAZIL: Piauí: Teresina: VIII/1852, A.K. Oliveira col., no host plant, 1 specimen (MNRJ). Rio Grande do Norte: Natal: III/1950, M. Alvarenga col., no host plant, 1 specimen (MNRJ). Mato Grosso: Nova Chavantina: Rio das Mortes, VII/1952, Alvarenga & Oliveira col., no host plant, 1 specimen (USNM). Porto Esperança: 2/II/1941, F. Lane col., no host plant, 2 specimens (DZUP). Bahia: no date, G. Bondar col., in leguminosae, 1 specimen (DZUP). Itaparica: 1/III/[19]91, C.S. Ribeiro-Costa col., no host plant, 17 specimens (DZUP). Distrito Federal: Brasília: VII/1960-[19]86, Exp. Formosa col., 1 specimen (MNRJ). Minas Gerais: Viçosa: 1/V/33, E.J. Hambleton col., no host plant, 1 specimen (USNM);10/VI/33, same collector and no host plant, 1 specimen (DZUP). Espírito Santo: Baixo Guandu: 23–30/IX/1971, C. Elias col., no host plant, 1 specimen (DZUP). Guarapari: IX/1960, M. Alvarenga col., no host plant, 1 specimen (DZUP). Mato Grosso do Sul: Campo Grande: 20/VIII/1987, no collector and host plant, 1 specimen (DZUP). Jardins: V-VI/2007, A.C.V. Pires et al. col., in Senna occidentalis, 5 specimens (DZUP). Rio de Janeiro: Arraial do Cabo: 10/X/1987, L.R.C. Souza col., in Senna bicapsularis, 3 specimens (DZUP). Barra de Maricá: III-VI/1996, Pimentel col., in Senna australis, 3 specimens (DZUP). Maricá: III-VI/1996, same collector and no host plant, 1 specimen (DZUP). Teresópolis: P.N. Serra dos Órgãos, VI/2006, J.H. Viana col., in Senna neglecta var. oligophylla, 1 specimen (DZUP); VII/2006, same collector and host plant, 1 specimen (DZUP); 05/VII/2006, same col-

Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 lector and host plant, 1 specimen (DZUP); 22/VIII/2006, same collector and host plant, 1 specimen (DZUP); 05/IX/2006, same collector and host plant, 1 specimen (DZUP). São Paulo: São Paulo: 10/XII/1973, V.N. Alin col., no host plant, 1 specimen (USNM). Campinas: 12–17/VII/1987, C.S. Ribeiro-Costa col., in Senna sp., 2 specimens (DZUP). Paraná: Cianorte: 14/X/1985, F. Giacomel col., in Senna sp., 11 specimens (DZUP). Arapongas: 22/VII/[20]03, A. Menezes Jr. col., in Senna occidentalis, 9 specimens (DZUP); 22/VII/[20]03, same collector, in Senna sp., 4 specimens (DZUP). Curitiba: XI/XII [no year], L.C. Haenle col., in Senna macranthera, 16 specimens (DZUP); 2/XI/1986, C.S. Ribeiro-Costa col., in Senna sp., 3 specimens (DZUP); Journal of Natural History 15

02/VI/1987, same collector, in Senna neglecta var. neglecta, 9 specimens (DZUP); IX/1987, C. Vargas col., in Senna sp., 33 specimens (DZUP); IX/1987, same collector and no host plant, 4 specimens (DZUP); 10/VIII/1988, C.S.Ribeiro-Costa col., in Senna sp., 20 specimens (DZUP); 22/XI/1995, same collector, in Senna multijuga, 1 specimen (DZUP); 25/X/1996, same collector, in Senna macranthera var. macranthera, 19 specimens (DZUP); XI/1996, same collector and host plant, 2 specimens (DZUP); Ed. Física – UFPR, 8/V/1997, same collector, in Senna neglecta var. neglecta, 1 specimen (DZUP); Ed. Física – UFPR, 13–18/VI/97, same collector and host plant, 3 specimens (DZUP); 4/VIII/1997, same collector and host plant, 1 specimen (DZUP); 6/VI/1997, same collector and host plant, 1 specimen (DZUP); 4/VIII/1997, same collector and host plant, 1 specimen (DZUP); 11/VII/1997, same collector and host plant, 1 specimen (DZUP); 18/VII/1997, same collector and host plant, 1 specimen (DZUP); 23/VII/1997, same collector and host plant, 1 specimen (DZUP); 13/VIII/1997, same collector and host plant, 3 specimens (DZUP); 22/VIII/1997, same collector and host plant, 3 specimens (DZUP); 15/V/[19]98, same collector and host plant, 6 specimens (DZUP); Rua Artur Loyola, XI-I (1998–1999), no collector, in Senna macranthera, 8 specimens (DZUP); 20/IX/2001, E. Caron col., no host plant, 1 specimen (DZUP); 28/IX/2001, same collector and no host plant, 7 specimens (DZUP); 08/X/2001, same collector and no host plant, 3 specimens (DZUP); 09/X/2001, same collector and no host plant, 1 specimen (DZUP); 10/X/2001, same collector and no host plant, 3 specimens (DZUP); 11/X/2001, same collector and no host plant, 9 specimens (DZUP); 12/X/2001, same collector and no host plant, 8 specimens (DZUP); 15/X/2001, same collector and no host plant, 7 specimens (DZUP); 16/X/2001, same collector and no host plant, 1 specimen (DZUP); 18/X/2001, same collector and no host plant, 3 specimens (DZUP); 19/X/2001, same collector and no host plant, 1 specimen (DZUP); 22/X/2001, same collector and no host plant, 5 specimens (DZUP); 23/X/2001, same collector and no host plant, 3 specimens (DZUP); 24/X/2001, same collector and no host plant, 1 specimen (DZUP); 25/X/2001, same collector and no host plant, 2 specimens (DZUP); 26/X/2001, same collector and no host plant, 6 specimens (DZUP); 27/X/2001, same collector and no host plant, 1 specimen (DZUP); 30/X/2001, same collector and no host plant, 1 specimen (DZUP); 05/XI/2001, same collector and no host plant, 2 specimens (DZUP); 06/XI/2001, same collector and no host plant, 1 specimen (DZUP); 07/XI/2001, same collector and no host plant, 1 specimen (DZUP); 18/XI/2001, same collector and no host plant, 2 specimens (DZUP); 24/XI/2001, same collector and no host plant, 1 specimen (DZUP); 26/XI/2001, same collector and no host plant, 2 specimens (DZUP); 15/VII/2008, C.S. Ribeiro-Costa col., in Senna neglecta var. neglecta, 13 specimens (DZUP). Guaratuba: Pontal do Itararé, 6/IV/2008, 1200 m, 25◦5254 S, 48◦5756 W, P.S. Grossi Col., no host plant, 2 specimens (DZUP). Ponta Grossa:

Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 29/VI/2000, A. Linzmeier col., in Senna macranthera, 16 specimens (DZUP); IAPAR, 29/VI/2000, C.S. Ribeiro-Costa col., in Senna macranthera, 47 specimens (DZUP).

Distribution Colombia (Valle del Cauca), Venezuela (Carabobo), Bolivia (Santa Cruz), Brazil (Bahia, Rio de Janeiro, Minas Gerais, São Paulo, Mato Grosso, Mato Grosso do Sul, Paraná). 16 J. Herzog Viana and C.S. Ribeiro-Costa

New records – Brazil (Rio Grande do Norte, Piauí, Distrito Federal, Espírito Santo).

Host plants Fabaceae: Caesalpinioideae: Senna pistaciifolia, S. pendula, S. occidentalis, S. bicapsularis, S. macranthera, S. multijuga, S. surattensis, S. splendida, S. alata. New record. Fabaceae: Caesalpinioideae: Senna neglecta var. oligophylla, S. neglecta var. neglecta, S. appendiculata (= S. australis). Note: Koller and Nunes (2006) found a few specimens of S. bondari in seed cap- sules of Memora peregrina (Bignoniaceae), however, it is unclear whether the larvae really consume its seeds. Hence, because of the scarce information of Koller and Nunes (2006) and the unusual Bignoniaceae as a host of Sennius, we will not consider it as a record for S. bondari until more information is added.

Discussion Sennius bondari differs from Sennius nappi mainly by black integument of elytron with white pubescence forming a semicircle at the midline, and a dense tuft of white pubescence on basal half of pygidium. Belongs to the abbreviatus group of Sennius established by Johnson and Kingsolver (1973) that has three subgroups: subgroup 1, with Sennius abbreviatus (Say 1824) and Sennius rufomaculatus (Motschoulsky 1874); subgroup 2 with Sennius medialis (Sharp, 1885) and Sennius durangensis Johnson and Kingsolver 1973; subgroup 3, with Sennius lebasi (Fähraeus 1839), Sennius leucostau- rus Johnson and Kingsolver 1973, and Sennius trinotaticollis (Pic 1930). It shares with these species the same male genitalia pattern, with internal sac trilobate, submedian region with small spicules, and large lateral clusters of spicules not aligned in subapical region at base of lateral lobes. Sennius bondari is more similar to the species of subgroup 3, in particular to Sennius leucostauros by sharing with it a horizontal band of white pubescence on elytra, and hinge sclerites of middle lobe larger than those seen in the other species of the subgroup. Sennius bondari has elytra with sparse golden pubescence and pygidium with dense macula of white pubescence in median basal region not extending to apex, whereas in Sennius leucostauros the golden pubescence is denser at the ﬁrst and second striae intervals, pygidium with dense white pubescence on median side extending to apex. In male genitalia, Sennius bondari has hinge sclerites wider than those of Sennius leucostauros, and the internal sac of Sennius bondari has clusters with large spicules aligned in the subapical region, as in Sennius leucostauros they are absent.

Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 Sennius margarete sp. nov. (Figure 4A–Q)

Description Dimension. Length (Pronotum–elytra) 2.46–3.32 mm. Width 1.52–2.08 mm.

Integument colour. Head, thorax, abdomen, pygidium and legs light brown to black (Figure 4A–L). Maxillary and labial palps dark brown. Hind legs and antennae light to dark brown. Anterior and middle legs light brown (Figure 4B, F, J). Journal of Natural History 17 Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013

Figure 4. Sennius margarete sp. nov. (A–D) Red pattern: (A) dorsal view; (B) lateral view; (C) head, frontal view; (D) pygidium; (E–H) stained pattern: (E) dorsal view; (F) lateral view; (G) head, frontal view; (H) pygidium; (I–L) black pattern: (I) dorsal view; (J) lateral view; (K) head, frontal view; (L) pygidium; (M) base of elytra striae; (N) part of hind leg, internal view; (O) hind tibia, external view; (P, Q) male genitalia: (P) median lobe with hinge sclerites and others sclerites in internal sac magniﬁed; (Q) lateral lobes. 18 J. Herzog Viana and C.S. Ribeiro-Costa

Pubescence. Head with dense tuft of light yellow pubescence posterior to postocular lobe, remainder of head with moderately dense pubescence. Labrum with scattered golden pubescence, slightly denser at apex (Figure 4C, G, K). Pronotum with moderately dense yellow and golden pubescence (Figure 4A, E, I). Scutellum with dense white pubescence (Figure 4A, E, I). Elytron intermixed with light yellow and golden pubescence homogeneously distributed, sometimes in moderately dense to dense tufts at base of third to seventh striae intervals (Figure 4A, E, I). Pygidium with moderately dense light yellow pubescence, often with denser tuft at middle-basal (Figure 4D, H, L). Ventral surface with light yellow pubescence denser at base of mesepimeron, posterior margin of metepisternum, and distal region of hind coxa (Figure 4B, F, J). Head with coarse punctuations except at midline of frons, with fine punctuations; labrum smooth; convex frons, frontal carina and transverse sulcus absent; ocular sinus deep, more than half eye length (Figure 4C, G, K). First and third antenna segments filiform, second and fourth moniliform, fifth to tenth wider than long, and eleventh globose with pointed apex (Figure 4B, F, J). Pronotum moderately convex, disc slightly sulcate at basal lobe (Figure 4A, E, I). Scutellum subquadrate, slightly wider than long (Figure 4A, E, I). Base of fourth and fifth striae of elytron with denticles (Figure 4M). Hind femur on ventral margin near the apex with very prominent tooth (0.06–0.1 mm) microserrate (Figure 4N). Hind tibia with lateroventral carina about half its length (Figure 4O), lateral, dorsomesal, and ventral carinae long, ending near the coronal denticles (Figure 4N, O); mucro about the same length as coronal denticles (Figure 4N, O). Last abdominal ventrite of male emarginate at middle, rounded in female. Pygidium not concealed by elytra, lateral margins slightly curved; apex rounded in male, apex more acuminate in female (Figure 4B, D, F, H, J, L).

Male genitalia. Median lobe about 5.4 times longer than its width at middle, expanded at apex, ventral valve triangular with lateral margins straight, apex slightly truncate; hinge sclerites long, slender, almost straight; internal sac trilobate, basal region near ventral valve with two small dense clusters of spicules, submedian region with dense spicules forming irregular sclerotized areas that extend along lateral sides, middle, and apical regions of the internal sac, more intense on lateral lobes (Figure 4P). Tegmen with lateral lobes separated by deep emargination, about 0.7 times the length of lateral lobes (Figure 4Q).

Type material Holotype, allotype and 10 paratypes deposited in DZUP with labels: “Brazil, RJ- Teresópolis | P.N. Serra dos Órgãos | VI/2006 | J.H. Viana col.” “ Em | Senna neglecta | Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 var. oligophylla”. Five paratypes with the same label as the holotype, deposited in each museum: MZSP, MNRJ, TAMU, USNM, FSCA, CEAM and CNCI.

Additional specimens BRAZIL: Rio de Janeiro: Teresópolis: P.N. Serra dos Órgãos, VI/2006, J.H. Viana col., in Senna neglecta var. oligophylla, 70 specimens (DZUP); 09/VI/2006, Journal of Natural History 19

same collector and host plant, 3 specimens (DZUP); 22/VIII/2006, same collector and host plant, 16 specimens (DZUP); 05/IX/2006, same collector and host plant, 11 specimens (DZUP); 02/V/2007, same collector and host plant, 3 specimens (DZUP); 02/VIII/2007, same collector and host plant, 5 specimens (DZUP); 15/VIII/2007, same collector and host plant, 1 specimen (DZUP); 17/VIII/2007, same collector and host plant, 1 specimen (DZUP); 22/VIII/2007, same collector and host plant, 3 specimens (DZUP); 29/VIII/2007, same collector and host plant, 1 specimen (DZUP). Paraná: Curitiba: 1997, Bobato col., in Senna neglecta var. neglecta, 1 specimen (DZUP); 12/VI/1997, C.S. Ribeiro-Costa col., no host plant, 17 specimens (DZUP); VI/1998, same collector, in Senna neglecta var. neglecta, 8 specimens (DZUP); 16/VI/1998, same collector and host plant, 4 specimens (DZUP); 15/VI/1998, same collector and host plant, 1 specimen (DZUP); Centro Politécnico, 29/II/2008, same collector and host plant, 8 specimens (DZUP); Ed Física – UFPR, 24/IV/2008, same collector and host plant, 2 specimens (DZUP); 27/VI/2008, same collector and host plant, 3 specimens (DZUP).

Distribution Brazil (Rio de Janeiro, Paraná).

Host plant Fabaceae: Caesalpinioideae: Senna neglecta var. oligophylla, S. neglecta var. neglecta.

Discussion Sennius margarete sp. nov. differs mainly by the pubescence of elytra usually evenly distributed, dense hair bands when present only at base (Figure 4A–L), and median lobe with nearly straight hinge sclerites (Figure 4P). Unlike Sennius bondari and Sennius nappi, which belong to the abbreviatus group, this species belongs to the fallax group of Sennius, composed of Sennius fallax (Boheman 1839), Sennius auricomus Johnson and Kingsolver 1973 (subgroup 1), Sennius alticola (Sharp, 1885), Sennius chalcodermus Johnson and Kingsolver 1973, Sennius discolor (Horn 1873), and Sennius atripectus Johnson and Kingsolver 1973 (subgroup 2), with particular resemblance to species of the last subgroup, sharing characters of the male genitalia, such as spicules distributed almost throughout the entire internal sac. Within subgroup 2 it is more similar to Sennius atripectus because both share the same distribution pattern of sclerites in internal sac, differing mainly by the shapes of the hinge sclerites (almost straight in

Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 Sennius margarete sp. nov., curved in Sennius atripectus) and shapes of the ventral valve (truncate at the apex in Sennius margarete sp. nov., rounded in Sennius atripectus).

Etymology The speciﬁc name “margarete” is in honour of Professor Margarete de Macedo Monteiro, for her great professional and personal contribution in instructing J.H. Viana. This is a noun in apposition, gender feminine. 20 J. Herzog Viana and C.S. Ribeiro-Costa

Sennius nappi Ribeiro-Costa and Reynaud 1998 (Figure 5A–I)

Sennius nappi Ribeiro-Costa and Reynaud 1998, p. 249 (original description, distribution, type, ﬁgures, host); Caron et al. 2004, p. 5 (egg description); Sari et al. 2005, pp. 169–174 (biology); Sari and Ribeiro-Costa 2005, pp. 521–525 (biology).

Diagnosis Dimension. Length (Pronotum–elytra) 1.80–2.84 mm. Width 1.08–1.72 mm.

Integument colour. Head black, labrum rarely lighter; ﬁrst four segments of antenna red-orange, seventh to apical dark brown to black (Figure 5B). Prothorax, scutellum, pygidium and ventral region of the body black (Figures 5A, B, D). Black elytron with transverse submedian macula, irregular, red-orange to dark brown, extending from third or fourth to eighth or ninth striae interval, sometimes extending more darkly to apex region, elytra rarely without macula (Figure 5A). Anterior and middle legs black or dark brown apically; black hind legs (Figure 5B). Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013

Figure 5. Sennius nappi. (A) Dorsal view; (B) lateral view; (C) head, frontal view; (D) pygidium; (E) base of elytra striae; (F) part of hind leg, internal view; (G) hind tibia, external view; (H, I) male genitalia: (H) median lobe with hinge sclerites and others sclerites in internal sac magniﬁed; (I) lateral lobes. Journal of Natural History 21

Pubescence. Head with dense tuft of white pubescence posterior to postocular lobe, moderately dense on postocular lobe and scattered on rest of head. Labrum with scattered golden pubescence, denser and forming a line near the apex (Figure 5C). Pronotum with brown and golden sparse pubescence and white pubescence denser in short oblique lateral bands, and usually in the antescutellar region (Figure 5A). Scutellum with dense white pubescence (Figure 5A). Elytron with moderately dense white pubescence forming a submedian transversal and irregular macula, extending from second or third to tenth striae interval, generally covering the orange-red macula on the integument; remainder of elytra with sparse brown pubescence (Figure 5A). Ventral surface with white pubescence denser at base of mesepimeron, at posterior margin of metepisternum, at distal region of hind coxa, and at base of first abdominal ventrite (Figure 5B). Pygidium with moderately dense white pubescence at the anterior midline, sometimes extending to the middle of pygidium (Figure 5D). Head with coarse and dense punctuation except at midline of frons and labrum; frons slightly convex, transverse sulcus slightly evident; ocular sinus deep, more than half an eye length (Figure 5C). First to fourth segments of the antenna moniliform to filiform, fifth to tenth wider than long, eleventh globular and pointed apically (Figure 5B). Pronotum moderately convex (Figure 5B), disc sulcate at basal lobe (Figure 5A). Scutellum subquadrate, slightly wider than long (Figure 5A). Base of second and third striae of elytron with denticles (Figure 5E). Hind femur on ventral margin with a slightly prominent microserrate tooth (0.01–0.06 mm) (Figure 5F). Hind tibia with lateroventral carina that extends more than half of its length (Figure 5G), lateral, dorsomesal and ventral carinae long, ending near coronal denticles (Figure 5F, G); mucro short, about 1.3 times the opposite coronal denticle (Figure 5F, G). Last abdominal ventrite of male emarginate at middle, rounded in female. Pygidium not concealed by elytra, subtriangular, lateral margins slightly curved, apex rounded in male, more acuminate in female (Figure 5B, D).

Male genitalia. Median lobe about ﬁve times longer than its width at median region, apex slightly expanded; ventral valve with rounded projection apically, lateral margins convex; hinge sclerites moderately curved, expanded at base and apex. Trilobate internal sac, without spicules at basal region near base of valve, submedian region with dense group of short spicules apparently forming three longitudinal clusters; laterals of subapical region of internal sac with two clusters of large spicules; apical region with spicules on lateral lobes and denticles on middle lobe near the gonopore (Figure 5H). Tegmen with lateral lobes separated by deep emargination, about 0.7 times the length of lateral lobes (Figure 5I).

Type material Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 Holotype, allotype and paratypes of Sennius nappi deposited in DZUP with the labels: “Curitiba-PR | 22-XI-95 | Ribeiro-Costa,” “Host | Senna | multijuga”. For the recognition of Sennius nappi, the type material of Sennius transversesignatus was studied.

Additional specimens BRAZIL: Mato Grosso: Pirizal: Faz. Retiro Novo, Pantanal de Poconé, N. Sra. do Livramento, 28/III/[20]04, L.D. Battirola col., in Vochysia divergens, 1 specimen 22 J. Herzog Viana and C.S. Ribeiro-Costa

(DZUP). Mato Grosso do Sul: Dourados: no date, A. Ferraz col., no host plant, 1 specimen (DZUP). Rio de Janeiro: Itatiaia: P.N. Itatiaia, 19/VIII/2008, M. Torres col., in Senna multijuga, 24 specimens (DZUP). Rio de Janeiro: X/1957, M. Alvarenga col., no host plant, 1 specimen (DZUP). Teresópolis: P.N. Serra dos Órgãos, VI/2006, J.H. Viana col., in Senna neglecta var. oligophylla, 4 specimens (DZUP); VIII/2006, same collector and host plant, 2 specimens (DZUP); 22/VIII/2006, same collector and host plant, 29 specimens (DZUP); 05/IX/2006, same collector and host plant, 18 specimens (DZUP); 21/IX/2006, same collector and host plant, 1 specimen (DZUP); 29/IX/2006, same collector and host plant, 2 specimens (DZUP); 2007, same collector and host plant, 2 specimens (DZUP); 02/IV/2007, same collector and host plant, 1 specimen (DZUP); 15/VIII/2007, same collector and host plant, 6 specimens (DZUP); 17/VIII/2007, same collector and host plant, 1 specimen (DZUP); 29/VIII/2007, same collector and host plant, 1 specimen (DZUP); 06/IX/2007, same collector and host plant, 1 specimen (DZUP); 12/IX/2007, same collector and host plant, 6 specimens (DZUP); 14/IX/2007, same collector and host plant, 4 specimens (DZUP); 21/IX/2007, same collector and host plant, 1 specimen (DZUP); 26/IX/2007, same collector and host plant, 1 specimen (DZUP); 26/IX/2007, same collector and host plant, 1 specimen (DZUP). Paraná: Curitiba: 22/XI/[19]95, C.S. Ribeiro-Costa col., in Senna multijuga, 9 specimens (DZUP); Jardim Botânico, 04/VIII/1997, same collector and host plant, 3 specimens (DZUP); 20/XI/[20]00, L.T. Sari col., same host plant, 3 specimens (DZUP); 26/I/[20]01, same collector and host plant, 3 specimens (DZUP); Ed. Física – UFPR, 15/VII/[20]08, K.L. Azevedo col., in Senna neglecta var. neglecta, 3 specimens (DZUP). Jaguariaíva: 03/X/[20]03, L.T. Sari col, in Senna rugosa, 1 specimen (DZUP); 24/X/[20]03, same collector and host plant, 1 specimen (DZUP); 21/X/[20]03, same collector and host plant, 1 specimen (DZUP). Londrina: 16/IV/2003, A. Menezes Jr. col., in Senna sp., 3 specimens (DZUP); 10/VI/2002, same collector, in Senna occidentalis, 2 specimens (DZUP); 22/VIII/2003, same collector, in Senna obtusifolia, 3 specimens (DZUP); 14/V/2002, same collector, in Senna sp., 3 specimens (DZUP). No locality: no date and collector, 1 specimen (DZUP).

Distribution Brazil (Paraná). New record – Brazil (Mato Grosso, Mato Grosso do Sul, Rio de Janeiro)

Host plants Fabaceae: Caesalpinioideae: Senna multijuga. New records – Fabaceae: Caesalpinioideae: Senna neglecta var. oligophylla, Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 S. neglecta var. neglecta, S. rugosa, S. occidentalis, S. obtusifolia. Note. The plant listed in the additional specimens, Vochysia divergens, is not listed here because the adult collected was not reared on seeds of this plant, and only plants with record of larvae preying are quoted as host plant.

Discussion Sennius nappi differs mainly by its pronotum with short oblique lateral bands of dense white pubescence, and elytron usually with an irregular transverse macula of Journal of Natural History 23

white pubescence in the submedian region covering a reddish-orange macula that may extend, becoming darker, to the apex of the elytron. Sennius bondari and Sennius nappi likewise belong to the abbreviatus group of Sennius by sharing many characters men- tioned in the “Discussion” section of Sennius bondari.However,Sennius nappi is placed in subgroup 1 and is more similar to Sennius transversesignatus, sharing two major clusters of spicules on each lateral lobe of the internal sac of the male genitalia (a characteristic of subgroup 1) and a black integument, except for a reddish-orange transverse macula on each elytron covered with white pubescence, which can make identiﬁcation difﬁcult. In Sennius nappi, the last seven antenna segments are dark brown to black, and there are no tufts of white pubescence near the base of the elytron, whereas in Sennius transversesignatus all segments are usually light brown and tufts of white pubescence are present. In terms of the male genitalia, Sennius nappi has lateral clusters of large spicules in the subapical region of the internal sac, whereas in Sennius transversesignatus these clusters are in the median region. Moreover, Sennius nappi does not have two moderately dense clusters of large spicules in the apical region of the middle lobe of the internal sac, as in Sennius transversesignatus.

Sennius ricardo sp. nov. (Figure 6A–H)

Description Dimension. Length (Pronotum-elytra) 2.44 mm. Width 1.60 mm.

Integument colour. Head, thorax and elytra black, elytron sometimes with transverse irregular dark brown macula in submedian region (Figure 6A) that extends from the third to eighth striae interval. Abdomen and pygidium dark brown to black (Figure 6B, D). Antennae, hind legs, maxillary and labial palps dark brown (Figure 6B, C). Fore and middle legs red-orange to dark brown (Figure 6B).

Pubescence. Head with dense tuft of white hair posterior to the postocular lobe, white pubescence moderately dense in the ocular sinus, postocular lobe and mandible (Figure 6C). Labrum with scattered golden pubescence, denser at apex (Figure 6C). Pronotum with sparse white and golden pubescence; white pubescence moderately dense at posterior corners and in antescutellar area, sometimes at anterior corners (Figure 6A). Scutellum with dense white pubescence (Figure 6A). Elytron with mix of black, white and golden pubescence, dense white pubescence distributed as follows: in small tufts at base of thirrd, fifth and sixth striae intervals; in submedian irregular transverse macula extending from first to ninth striae intervals, generally covering the Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 macula with a dark brown integument (Figure 6A). Pygidium with sparse pubescence on anterior half, and with moderately dense white pubescence and dense median line on posterior half (Figure 6D). Ventral region of body with white pubescence, denser at base of mesepimeron, metepisternum, metasternum, distal region of posterior coxa, and base of first abdominal ventrite (Figure 6B). Head evenly punctuate, except at midline of frons, base of clypeus and labrum; frons convex, frontal carina and transverse sulcus absent; ocular sinus deep, more than half an eye length (Figure 6C). First and third antenna segments filiform, second and fourth moniliform, fifth to tenth broader than long, eleventh globular and pointed at 24 J. Herzog Viana and C.S. Ribeiro-Costa

Figure 6. Sennius ricardo sp. nov. (A) Dorsal view; (B) lateral view; (C) head, frontal view; (D) pygidium; (E) part of hind leg, internal view; (F) hind tibia, external view; (G, H) male genitalia: (G) median lobe with hinge sclerites and others sclerites in internal sac magniﬁed; (H) lateral lobes.

apex (Figure 6B). Pronotum moderately convex, disc sulcate at basal lobe (Figure 6A). Scutellum subquadrate, slightly wider than long (Figure 6A). Base of elytra without denticles at striae. Hind femur with a very prominent tooth (0.05 mm), microserrate on ventral margin near apex (Figure 6E). Hind tibia with lateroventral carina extending more than half its length (Figure 6F); lateral, dorsomesal and ventral carinae long, ending near coronal denticles (Figure 6E, F); mucro slightly larger than the opposite coronal denticle (Figure 6E, F). Last abdominal ventrite of male emarginate at middle, rounded in female. Pygidium not concealed by elytra, lateral margins slightly curved, rounded apex in male, more acuminate in female (Figure 6B, D).

Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 Male genitalia. Median lobe about 4.3 times longer than its width at middle, not expanded at apex; ventral valve triangular with lateral margins straight and slightly rounded at apex; hinge sclerites of median lobe short; internal sac at basal region without spicules near base of ventral valve, submedian region with dense cluster of short spicules evenly distributed, apparently forming two more sclerotized longitudinal masses; subapical region with denticles and tiny microserrate sclerites; spicules slightly larger than those of submedian region and slightly denser than at lateral lobes of internal sac (Figure 6G). Tegmen with lateral lobes separated by deep emargination, about 0.7 times the length of lateral lobes (Figure 6H). Journal of Natural History 25

Distribution Brazil (Rio de Janeiro).

Host plant Fabaceae: Caesalpinioideae: Senna neglecta var. oligophylla.

Discussion Sennius ricardo sp. nov. differs mainly by having pronotum with dense white pubescence on posterior corners and antescutellar area, as well as small tufts at base of third, ﬁfth and sixth striae intervals of the elytron (Figure 6A), hind femur with very prominent microserrate tooth (Figure 6E), and hind tibia with lateroventral carina longer than half its length (Figure 6F). In terms of the male genitalia, the hinge sclerites are short, and the internal sac has denticles and is microserrate, with tiny sclerites at the subapical region and spicules slightly larger than those of submedian region and slightly denser than those of the lateral lobes (Figure 6G). As in the previous species, Sennius ricardo sp. nov. shows a pattern of male genitalia similar to that found in species of subgroup 2 of the fallax group of Sennius. It is similar to Sennius atripectus by its male genitalia pattern, but in Sennius ricardo sp. nov. the basal region has no spicules near the base of the valve (present in Sennius atripectus) and hinge sclerites are more curved in this species and less curved in Sennius atripectus.

Etymology The speciﬁc name “ricardo” is in honour of Professor Ricardo Ferreira Monteiro, for his great professional and personal contribution to instructing J.H. Viana. This is a

Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 noun in apposition, gender masculine.

Final considerations There have been only two previous papers from Brazil focusing on bruchines associated with host plants that provided identiﬁcation keys and species illustrations, including male genitalia – one on the beetles hosted by Senna multijulga and the other on Chamaecrista species (Ribeiro-Costa and Reynaud 1998; Silva et al. 2003). We focused on Senna neglecta in this paper, and for the ﬁrst time recorded it as a 26 J. Herzog Viana and C.S. Ribeiro-Costa

host for six species of Bruchinae, three of them new from three associated genera. Five species belonged to Bruchini, the largest tribe in the subfamily (with 80% of the known species); only one species belonged to Amblycerini, also a very diverse tribe. Among the species of Bruchini, Acanthoscelides multimaculatus sp. nov. is allocated to the group quadridentatus of Acanthoscelides that now comprises 16 species. Sennius bondari and Sennius nappi, also of Bruchini, were placed in the abbreviatus group, increasing the number of species in this group to nine. Sennius margarete sp. nov. and Sennius. ricardo sp. nov. were included here in the fallax group, subgroup 2, which now has eight species. Only one species of Acanthoscelides and one of Amblycerus, the richest genera of Bruchinae, were recorded consuming S. neglecta seeds (Acanthoscelides multimaculatus sp. nov. and Amblycerus hoffmanseggi). Sennius had the highest number of species feeding on this host – four species; two of them are described here, resulting in a total of 59 species now assigned to this genus. A possible explanation for this high number of Sennius species associated with Senna neglecta is the special preference of this genus for Cassiinae hosts, particularly Senna. Among all of the species recorded in association with Senna neglecta, Sennius bondari is the most polyphagous, for it has been found associated with a total of 11 Senna species (and even one different plant family). The interactions between bruchines and their host plants are subject to variables such as temporal, seasonal and spatial changes, parasitism and competition, making it unlikely to ﬁnd all of these seed predator species at the same time in the same place (Romero et al. 2009). Long-term studies with many types of fruits and seeds throughout the different host plants’ reproductive cycles will be necessary to gather complete records of all host associations and to survey all the species in speciﬁc localities.

Acknowledgements We are grateful to the curators listed in the Material and Methods section for loans; Dr S.P. Rosa (USP) for the type images from BMNH; TAXon line – Rede Paranaense de Coleções Biológicas (UFPR) for the photographs; and the Conselho Nacional de Desenvolvimento Cientíﬁco e Tecnológico (CNPq) for the scolarships to the authors. Contribution number 1856, Departmento de Zoologia, Universidade Federal do Paraná, Brazil.

References Arora GL. 1977. Taxonomy of the Bruchidae (Coleoptera) of Northwest India. Delhi: Association for the Study of Oriental Insects; p. 114–115. Blackwelder RE. 1946. Checklist of the coleopterous insects of Mexico, Central America, the

Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 West Indies, and South America. Bull U S Natl Mus. 185:551–763. Blair KG. 1928. Coleoptera (Heteromera, Teredilia, Malacodermata and Bruchidae) from the Galapagos Islands, collected on the “St. George” Expedition, 1924. Ann Mag Nat Hist. 1:671–680. Bondar G. 1931. Notas biológicas sobre bruquídeos brasileiros do gênero Spermophagus. Correio Agrícola. 11:55–59. Bondar G. 1936. Notas biológicas sobre bruquídeos observados no Brasil. Arch Inst Biol Vegetal. 3:7–44. Borowiec L. 1987. The genera of seed-beetles (Coleoptera, Bruchidae). Pol Pismo Entomol. 57:3–207. Journal of Natural History 27

Bosq JM. 1943. Segunda lista de Coleópteros de la República Argentina, dañinos a la Agricultura. Ing Agron. 4:44–47. Bridwell JC. 1944. A new Amblycerus affecting seeds of Prosopis chilensis in Puerto Rico and Hispaniola. J Agric Univ P R. 27:133–135. Caron E, Ribeiro-Costa CS, Linzmeier AM. 2004. The egg morphology of some species of Sennius Bridwell (Coleoptera: Chrysomelidae: Bruchinae) based on scanning electron micrographs. Zootaxa. 556:1–10. Casari A, Teixeira EP. 1997. Descriptions and biological notes of final larval instar and pupa of some seed beetles (Coleoptera: Bruchidae). Ann Soc Entomol Fr. 33:295–321. Costa Lima A. 1928. Segundo catálogo systemático dos insectos que vivem nas plantas do Brasil e ensaio de bibliographia entomológica brasileira. Arch Esc Super Agric Med Vet. 8:69–301. Costa Lima A. 1955. Insetos do Brasil. Coleópteros, Tomo 9, 3a parte. Rio de Janeiro: Escola Nacional de Agronomia, Série Didática; p. 289p. Dejean PFMA. 1837. Catalogue des Coléoptères de la collection de M. Le Comte Dejean. 3a ed., v. 5. Paris: Chez Méquignon-Marvis Pére et Fils; p. 384p. Delobel A, Deloel H, Tran M, Sembène M, Han SH. 1995. Observations sur les relations trophiques entre les bruches du genre Caryedon (Coléoptères, Bruchidae) et leurs plantes hôtes sauvages au Sénégal. Bull. Inst. fond. Afi Noire Cheikh Anta Diop, Sér A. 48:78–88. Fabricius JC. 1781. Species insectorum. Kilonii. 1:552p. Gemminger M, Harold B. 1873. Catalogus coleopterorum hucusque descriptorum synonymicus et systematicus [Cerambycidae (Lamiinae), Bruchidae]. v. 10, pars I, Paris: p. 2989–3232. Guérin J. 1953. Coleópteros do Brasil. São Paulo: Faculdade de Filosofia, Ciências e Letras da Universidade de São Paulo, Departamento Zoologia, Fisiologia Geral e Animal; p. 356p. Gyllenhal L. 1833. In: Schoenherr CJ, Genera et species, curculionidum, cum synonymia hujus familiae. Volume 1. Paris; 381p. ILDIS 2005 ILDIS: International Legume Database & Information Service [Internet]. Version 10, November 2005. Reading (Berkshire): Centre for Plant Diversity and Systematics; [cited 2010 April]. Available from: http://www.ildis.org/. Irwin HS, Barneby RC. 1982. The American Cassiinae. Mem N Y Bot Garden. 35:1–918. Jekel H. 1855. Insecta Saundersiana: or characters of undescribed insects in the collection of William Wilson Saunders. (Coleoptera) (Bruchidae 1-37), London; 242p. Johnson CD. 1984. Sennius yucatan, new species, a redescription of S. infractus, and new host records for other Sennius (Coleoptera: Bruchidae). Ann Entomol Soc Am. 77:56–64. Johnson CD. 1989. Adaptive Radiation of Acanthoscelides in Seeds: Examples of Legume- Bruchid Interactions. In: Stirton CH, Zarucchi JL, editor. Advances in Legume Biology. Monographs in Systematic Botany from the Missouri Botanical Garden, Vol. 29. St. Louis, MO; p. 747–779. Johnson CD. 1990. Systematics of the seed beetle Genus Acanthoscelides (Bruchidae) of Northern South America. Trans Am Entomol Soc. 116:297–618. Johnson CD, Kingsolver JM. 1973. A Revision of the genus Sennius of North and Central America (Coleoptera: Bruchidae). U.S. Department of Agriculture Technical Bulletin. 135p. Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 Johnson CD, Kingsolver JM. 1982. Checklist of the Bruchidae (Coleoptera) of Canada, United States, Mexico, Central America, and the West Indies. Coleopt Bull. 35:409–422 [1981). Johnson CD, Romero Nápoles J, Raimúndez-Urrutia E. 2001. Ecology of Amblycerus cras- sipunctatus Ribeiro-Costa (Coleoptera: Bruchidae) in seeds of Humiriaceae, a new host family for bruchids, with an ecological comparison to other species of Amblycerus. Coleopt Bull. 55:37–48. Kingsolver JM. 1970. A synopsis of the subfamily Amblycerinae Bridwell in the West Indies, with descriptions of new species (Coleoptera: Bruchidae). Trans Am Entomol Soc. 96:469–497. 28 J. Herzog Viana and C.S. Ribeiro-Costa

Kingsolver JM. 1971. Description of new seed beetle from Australia (Coleoptera: Bruchidae). J Aust Entomol Soc. 10:179–182. Kingsolver JM. 1980. The Quadridentatus group of Acanthoscelides: descriptions of three new species, notes, synonymies, and a new name (Coleoptera, Bruchidae). Brenesia. 17:281–294. Kingsolver JM. 2004. Handbook of the Bruchidae of the United States and Canada (Insecta, Coleoptera), Vol. 1. Technical Bulletin No. 1912. Washington, (DC): United States Department of Agriculture; p. 1–324. Kingsolver JM, Silva P. 1991. Update of scientific names of Bruchidae (Coleoptera) listed by Bondar in “Notas Biológicas” (1931 and 1936). An Soc Entomol Brasil. 20:411–415. Koller WW, Nunes SG. 2006. Proposta de Manejo de Memora peregrine – a Ciganinha. Campo Grande: Embrapa Gado de Corte, Documentos, 101; p. 9p. Leal L, Biondi D. 2006. Influência de substratos e tratamentos pré-germinativos em sementes de Senna neglecta (Vogel) H.S. Irwin & Barneby. In: 57◦ Congresso Nacional de Botânica / 13◦ Encontro Estadual de Botânicos / 5◦ Encontro Estadual de Herbários, Gramado. Sociedade Botânica do Brasil; p. 646–646. Leal L, Biondi D, Guimarães O, Santos JB Jr. 2004. Estudos preliminares de espécies nativas para o uso no paisagismo. In: 12◦ EVINCI – Evento de Iniciação Científica, UFPR – Curitiba – PR. Livro de Resumos; p. 240. Leng CW. 1920. Catalogue of the Coleoptera of America, north of Mexico. New York: John D. Sherman Jr; p. 470p. Linzmeier AM, Ribeiro-Costa CS, Caron E. 2004. Comportamento e ciclo de vida de Sennius bondari (Pic, 1929) (Coleoptera, Chrysomelidae, Bruchinae) em Senna macranthera (Collad.) Irwin et Barn. (Caesalpinaceae). Rev Bras Zool. 21:351–356. Macêdo MV, Lewinsohn T, Kingsolver JM. 1992. New host records of some bruchid species in Brazil with the description of a new species of Caryedes (Coleoptera: Bruchidae). Coleopt Bull. 46:330–336. Marazzi B, Endress PK, Queiroz LP, Conti E. 2006. Phylogenetic relationships within Senna (Leguminosae, Cassiinae) based on three chloroplast DNA regions: patterns in the evolution of floral symmetry and extrafloral nectarines. Am J Bot. 93:288–303. Olivier AG. 1795. Entomologie, ou histoire naturelle des insects. Coléoptères. v. 4, Lanneau, Paris; 79p. Pic M. 1913. In: Junk W. Coleopterorum Catalogus. Pars 55, Bruchidae, Berlim v,26., 74p. Pic M. 1929. Nouveautes diverses. Melanges Exotico Entomol. 53:1–36. Pierce D. 1915. Descriptions of some weevils reared from cotton in Peru. J Agric Res. 102:1–16. Pimentel M. 1997. Interações ecológicas envolvendo duas espécies de Sennius (Coleoptera: Bruchidae) predadores de sementes de sua planta hospedeira Senna australis (Leguminosae). Dissertação de Mestrado. Universidade Federal do Rio de Janeiro, Rio de Janeiro; 59p. Ribeiro-Costa CS. 1992. Gênero Amblycerus Thunberg, 1815 (Coleoptera: Bruchidae). Grupo hoffmanseggi: II. Redescrições, chave e dados biológicos das espécies. Rev Bras Entomol. 36:149–175. Ribeiro-Costa CS. 1998. Observations on the biology of Amblycerus submaculatus (Pic) and Sennius bondari (Pic) (Coleoptera: Bruchidae) in Senna alata (L.) Roxburgh Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013 (Caesalpinaceae). Coleopt Bull. 52:63–69. Ribeiro-Costa CS, Marinoni RC. 1992. Gênero Amblycerus Thunberg, 1815 (Coleoptera: Bruchidae). Grupo “hoffmanseggi”: I. Taxonomia Numérica. Rev Bras Entomol. 36:149–175. Ribeiro-Costa CS, Reynaud DT. 1998. Bruchids from Senna multijuga (Rich) I. & B. (Caesalpinaceae) in Brazil with descriptions of two new species. Coleopt Bull. 52: 245–252. Rodrigues RS, Flores AS, Miotto STS, Baptista LRM. 2005. O gênero Senna (Leguminosae, Caesalpinioideae) no Rio Grande do Sul, Brasil. Acta Bot Brasilica. 19:1–16. Journal of Natural History 29

Romero J, Johnson CD. 2000. Revision of the genus Zabrotes Horn of Mexico (Coleoptera: Bruchidae: Amblycerinae). Trans Am Entomol Soc. 126:221–274. Romero Nápoles J, Johnson CD. 2002. BRUCOL: a data base for Bruchidae (Insecta: Coleoptera). In: Romero JN, Estrada, EV, Equihua AM, editors. Entomología Mexicana. Sociedad Mexicana de Entomología, vol. 1. México; p. 520–524. Romero Nápoles J, Johnson CD. 2004. Zabrotes maesi, A New Species from Nicaragua with New Distribution Records for other Bruchids (Coleoptera: Bruchidae: Amblycerinae). Coleopt Bull. 58:379–387. Romero Nápoles J, Perez AC, Kingsolver JM. 2009. Seed beetles (Coleoptera: Bruchidae) associated with Acacia cornigera (L.) Willd., with description of a new species of Acanthoscelides Schilsky. Insecta Mundi. 93:1–11. Sari LT, Ribeiro-Costa CS. 2005. Predação de sementes de Senna multijuga (Rich.) H. S. Irwin & Barneby (Caesalpinaceae) por bruquíneos (Coleoptera: Chrysomelidae). Neotrop Entomol. 34:521–525. Sari LT, Ribeiro-Costa CS, Roper JJ. 2005. Dinâmica populacional de bruquíneos (Coleoptera, Chrysomelidae) em Senna multijuga (Rich.) H. S. Irwin & Barneby (Caesalpinaceae). Rev Bras Zool. 22:169–174. Sharp D. 1885. Biologia Centrali-Americana. Insecta Coleoptera Bruchidae. 5:437–504. Silva AGDA, Gonçalves CR, Galvão DM, Gonçalves AJL, Gomes J, Silva NM, Simoni L. 1968. Quarto catálogo dos insetos que vivem nas plantas do Brasil, seus parasitos e predadores. 1◦tomo. Insetos, hospedeiros e inimigos naturais. Parte II. Ministério da Agricultura, Laboratório Central de Patologia Vegetal. Rio de Janeiro; 622p. Silva JAP, Ribeiro-Costa CS, Johnson CD. 2003. Sennius Bridwell (Coleoptera, Bruchidae): novas espécies predadoras de sementes de Chamaecrista Moench (Caesalpinaceae) da Serra do Cipó, Santana do Riacho, Minas Gerais, Brasil. Rev Bras Zool. 20:269–277. Souza GC, Haas APS, Von Poser GL, Schapoval EES, Elisabetsky E. 2004. Ethnopharmacological studies of antimicrobial remedies in the south of Brazil. J Ethnopharmacol. 90:135–143. Terán A, L‘Argentier SM. 1981. Observaciones sobre Bruchidae (Coleoptera) del noroeste argentino. IV. Estudios morfológicos y biológicos de Amblycerus hoffmanseggi (Gyll.), Acanthoscelides computus Kingsolver y Merobruchus bicoloripes (Pic). Acta Zool Lilloana. 36:61–84. Udayagiri S, Wadhi SR. 1989. Catalog of Bruchidae. Mem Am Entomol Inst. 45:301p. Zacher F. 1952. Die Nahrpﬂanzen der Samenkäfer. Z Angew Entomol. 33:460–480. Downloaded by [University Federal Do Parana], [Jéssica Viana] at 10:18 06 August 2013