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Systena Chevrolat 16 July 1998 PROC. ENTOMOL. SOC. WASH. 100(3), 1998, pp. 467-483 SYSTENA CHEVROLAT (COLEOPTERA: CHRYSOMELIDAE: ALTICINAE): NOTES ON NOMENCLATURE, REDESCRIPTION OF THE GENUS, AND A PRELIMINARY DISCUSSION OF CHARACTERS AND PHYLOGENETIC RELATIONSHIPS STEVEN W. LINGAFELTER, ALEXANDER S. KONSTANTINOV, AND JONG EUN LEE (SWL, ASK) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, c/o National Museum of Natural History, MRC-168, Washington, DC 20560 U.S.A. (email: [email protected]; akonstanti­ [email protected]); (JEL) Department of Biology, College of Natural Sciences, An­ dong National University, Andong, Kyungbuk 760-749, South Korea. Abstract.-A summary of the taxonomic history for Systena Chevrolat (Coleoptera: Chrysomelidae: Alticinae) is presented. Detailed discussion and illustration of morpho­ logical characters are presented based on stereo, compound, and scanning electron mi­ croscopy of dissected exemplar specimens for eight species of Systena from North, Central, and South America. A discussion of putative relationships of Systena to other alticine genera is presented. Key Words: Systena, Alticinae, Chrysomelidae, adult, morphology, character, nomencla­ ture, systematic, flea beetle, leaf beetle To help provide a foundation for future TAXONOMIC HISTORY generic-level phylogenetic studies in Al­ Systena was proposed by Chevrolat ticinae, a nomenclatural discussion and (1836:390). Heikertinger and Csiki (1939) detailed morphological redescription with and Blackwelder (1946) incorrectly attrib­ preliminary phylogenetic discussion are ute authorship of Systena to Melsheimer provided for Systena. Although the genus (1847), who indicated Chevrolat as author Systena was proposed in 1836, the first of Systena. In Melsheimer (1847), one ad­ discussion of morphological characters of ditional species, Systena blanda, was de­ the genus was presented by Clark (1865). scribed, but no other species were listed. However, as is typical of many early Chevrolat (1848) stated that Systena was works, the characters he used were super­ proposed by "us" (meaning himself) and ficial and not extremely useful consider­ adopted by Dejean. In this work, Systena ing the paucity of knowledge of character s-littera (Linnaeus) and S. vittata (Fabri­ distribution among Alticinae on a world­ cius) were mentioned as additional mem­ wide level. When characters are viewed in bers of the genus. Clark (1865) in his ex­ this context, as we aspire to do here, their amination of South American Alticinae pro­ value as diagnostic and evolutionary in­ vided the first characterization of Systena, dicators is greatly enhanced. and probably for this reason, Gemminger 468 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Character state information for the eight Systena species used for the generic characterization. Character numbers refer to text. Symbol, '' +'' indicates that both states present or too difficult to interpret. 'illTaxa Characters-+ 5 10 15 20 23 Systena blanda (Melsheimer) 00101 00030 010+0 +ODDO 000 Systena carri Blake 00001 00020 01000 10??0 ??? Systena championi Jacoby 00001 01020 01000 00010 000 Systena elongata Fabricius 00000 00020 01000 00010 000 Systena frontalis (Fabricius) 00001 00030 01000 00110 000 Systena marginallis (Illiger) 00001 00020 01000 00000 000 Systena oberthuri Baly +0001 00030 01000 +0000 001 Systena s-littera (Linnaeus) 00001 00020 01000 +ODDO 000 and von Harold (1874), Hatch (1927), Hom erthuri Baly, and S. s-littera (Linnaeus). (1889), and Leng (1920) considered him to These taxa which occur in North, Central, be the author of the genus. However, the and South America are suggested to repre­ mere use of one or more available species­ sent the spectrum of morphological diver­ group names in combination or clearly in­ sity for Systena. Characters, many of which cluded under a new genus-group name (as are described for the first time in Chryso­ done by Chevrolat 1836), is sufficient for melidae, are discussed in detail below. A availability for names published before subset of the characters and states is coded 1931 (ICZN [1985] Article 12 b-5; sum­ in Table 1 to explicitly indicate variability marized in Barber and Bridwell 1940). Oth­ or constancy within Systena. The appendix er authors who also correctly attribute to provides a brief description of the charac­ Chevrolat authorship of Systena include ters and states used in Table 1. Chapuis (1875), Crotch (1873), Seeno and Characters of adult exemplars were ex­ Wilcox (1982), and Poole and Gentili amined using a Leica MZ-APO stereo­ (1996). Systena frontalis (Fabricius) was scope, Leitz Diaplan compound microscope designated as the type species by Monr6s fitted with transmitted light interference and Bechyne (1956). contrast apparati (ICT), and an AMRAY 1810 scanning electron microscope. To ease METHODS examination of sclerotized features, adult The discussion of the adult characters specimens were partially disarticulated and and terminology follows that of Konstanti­ placed in hot KOH for several seconds. Ste­ nov and Lopatin ( 1987), Konstantinov reo and compound microscopic illustrations (1994) and Konstantinov and Vandenberg were made using camera lucidas. Images (1996). Hind wing terminology follows Ku­ were scanned at 600 dpi using a Nikon kalova-Peck and Lawrence (1993, Fig. 36) Scantouch scanner and enhanced using since they provided the most explicit hy­ Adobe Photoshop 3.0 and labeled using potheses of homology based on examina­ Macromedia Freehand 5.5 on a Power Mac­ tion of all beetle families. Metafemoral intosh 8500/132 with 50 megabytes RAM. spring terminology follows Furth (1982). Many characters are described here for the REDESCRIPTION OF SYSTENA CHEVROLAT first time. Morphological characters are Body (Fig. 1): Moderately elongate, flat based on at least two examples of dissected, in lateral view. Body length/width ratio as disarticulated specimens of Systena blanda measured by length of elytra divided by (Melsheimer), S. carri Blake, S. championi greatest width, ranging from 1.42 (S. s-lit­ Jacoby, S. elongata Fabricius, S. frontalis tera)-1.91 (S. oberthuri). Elytron without (Fabricius), S. marginallis (Illiger), S. ob- apical declivity. VOLUME 100, NUMBER 3 469 Fig. 1. Systena blanda, male, dorsal view. Head (Figs. 1, 7, 8): Antenna! calli pres­ tinct on anterior margin of antenna! calli but ent, well developed, each as large as anten­ otherwise shallow and not differentiated na! socket, longer than wide, extending to from surrounding area; not continuing be­ antenna! sockets. Supra-antennal sulcus dis- yond antenna! socket. Supraorbital sulcus 470 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON shallow, not well-differentiated from sur­ constricted at apical two-thirds, becoming rounding area. Supracallinal sulcus varies cone-shaped apically. Second antennomere from well developed in S. frontalis to poor­ shorter than third, longer than fourth. Fifth ly developed in S. blanda. Anterofrontal antennomere longer than fourth and sixth. ridge with distinct, narrow crest, lower than Mesal margins of antennal sockets farther frontal ridge, contacting frontal ridge an­ apart than diameter of antennal socket (Fig. teapically (before anterior end). Orbital sul­ 7). cus absent (when present in other genera, Thorax: Pronotum (Fig. 1) without pos­ extends from dorsal end of the supraorbital terior (prebasal) transverse impression and sulcus to dorsal margin of eye); dorsal mar­ without postero-lateral longitudinal impres­ gin of eye not demarcated from adjacent sions. Anterolateral and posterolateral cor­ vertex. Frontal ridge well developed, ex­ ners of pronotum each bearing one long tending between antenna! calli to clypeus seta. Procoxal cavities closed (apex of in­ (declivous between antennal calli), not tercoxal prostemal process contacting hy­ higher than rim of antennal socket when pomeral projection). Intercoxal prostemal viewed from antero-ventral perspective process strongly expanded apically (at least along plane of labrum. Frontal-lateral sul­ 2.0 times as wide at apex as narrowest re­ cus poorly developed; region between fron­ gion between procoxae). Mesocoxa (Fig. tal ridge and antennal socket not well dif­ 32) broadly open laterally to mesepimeron ferentiated. Subantennal sulcus absent. Sub­ (mesostemum and metastemum not con­ genal suture well-developed. Tentorium in­ tacting one another on the outer lateral mar­ complete, with only posterior arms gin of the mesocoxa). Intercoxal mesoster­ extending dorsally less than one-fifth of dis­ nal process (Fig. 32) bilobed. Mesonotum tance from floor of the head to the vertex. (Fig. 34) strongly sclerotized and lacking Occipital opening nearly evenly oval, not stridulatory ridges. Mesoprescutum (scutel­ subquadrate. lum, Fig. 34) is generally triangular, but Mouthparts (Figs. 2-5): Labrum (Fig. 2) more truncate posteriorly in S. blanda and with submarginal row of fine sensilla on in­ S. s-littera. Metendosternite (Figs. 37, 38) ner (ventral) surface; four long sensilla on with apex of anterior arms defiexed and ta­ outer (dorsal) surface; two short setae on pering to a point; with long, narrow, dorsal mid-apical margin on inner (ventral) sur­ ridge from anterior midpoint to posterior face; and with erratic pattern of short, pitch­ midpoint (Fig. 37); with poorly developed er-shaped sensilla on inner (ventral) sur­ tendons of metafurcal-mesofurcal muscles; face. Prementum (Fig. 3) with four setae with poorly developed
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