US ISSN 0006-9698

CAMBRIDGE,MASS.3FEBRUARY 2015 NUMBER 541

WHAT IS THE LEAF CALLIGRAPHA SCALARIS (LECONTE)?

JESU´ S GO´ MEZ-ZURITA1

ABSTRACT. Since the description of Calligrapha scalaris (LeConte, 1824), a reported as an occasional serious pest of American elm, this taxon has been affected by remarkable taxonomic confusion. Most authors have invoked their particular and generally flawed concept of C. scalaris (LeConte) without reference to John E. LeConte’s type material at the Museum of Comparative Zoology (Cambridge, Massachusetts). In this study, I have revised the series for C. scalaris (LeConte) conserved as part of J. LeConte and G. Horn collections, and confirmed that it includes as many as nine Calligrapha species, most of them lacking an association with Ulmus. Among these specimens, a couple of them from Texas are recognized as generally consistent with the original description and the only potentially elm-feeding . Thus, despite legitimate doubts for their availability to J. LeConte at the moment of the species description, they are designated here as neo- and paraneotypes, respectively, in an effort to maximize taxonomic stability. The species is redescribed on the basis of the neotype and diagnosed from all other species in this group, defined here as the ‘‘Calligrapha scalaris’’ group, including a provisional identification key for 14 species and for specimens conforming to the respective types. Finally, the study of syntype material from the Museum fu¨r naturkunde (Berlin) of C. multiguttis (Sta˚l, 1865), an early synonym of C. scalaris (LeConte), allowed recognition of several other Calligrapha taxa affected by this synonymy, including C. ignota Brown, C. knabi Brown, and C. tiliae Brown (5 Chrysomela multiguttis Sta˚l [pars] nov. syn.).

INTRODUCTION Go´mez-Zurita, 2013, 2014), no other species has been more problematic for its identifica- Since I started to study the biology, tion in collections than C. scalaris (LeConte, evolution, and, more recently, systematics 1824). Every author in every collection seems of the leaf beetle genus Calligrapha Chevro- to interpret this taxon in a slightly different lat, 1836 (Go´mez-Zurita, 2005; Go´mez-Zur- way, ‘‘a matter of individual opinion’’, as ita et al., 2004, 2006; Montelongo and pointed out by Knab (1909). The difficulties associated with this taxon were nicely illus- 1 Biodiversity and Evolution, Institut de Biologia Evolutiva (CSIC-Univ. Pompeu Fabra), Pg. trated by W. J. Brown (1945), who demon- Marı´tim Barceloneta 37, 08003 Barcelona, Spain; strated the existence of a species complex of e-mail: [email protected] very similar entities, at least morphologically,

E The President and Fellows of Harvard College 2015. 2 BREVIORA No. 541

Figure 2. Elytral spot nomenclature in a typical Figure 1. a, John Eatton LeConte’s artistic rendition species of the Calligrapha scalaris (LeConte) group of the species Chrysomela scalaris in the original 1824 (redrawn from figure 67 in Wilcox, 1972). publication, and b, a mounted specimen of Calligrapha scalaris (LeConte) collected on Ulmus in Oklahoma, SW of Crawford, Spring Creek Lake, 24 June 2011, T. ones that would be treated as Calligrapha in Montelongo leg. (IBE-JGZ voucher no. C326). the future. However, as the Nearctic fauna was being scrutinized, new species were that nevertheless differed strikingly in their described belonging to this group of animals ecologies, and even in their reproductive and soon the ‘‘diagnostic’’ characters for biologies, for some species are thelytokous. Calligrapha scalaris proved insufficient, re- This author described several new species sulting in the remarkable taxonomic confu- around C. scalaris and tried to provide sion occurring today for this species. Indeed, morphological arguments for their separation, LeConte’s characters correspond respective- yet the chief character to distinguish them ly, in the nomenclature of Wilcox (1972), to remained almost exclusively the knowledge confluent humeral spot and lunule, and to about their host plant (Brown, 1945; Wilcox, sutural and subsutural stripes fused together 1972). Unfortunately, this information is not with the arcuate band (Fig. 2). These char- available for most collection specimens, which acters are shared by at least 13 currently accepted North American taxa (and many therefore remain unidentifiable, at least until southern Nearctic and Neotropical species, diagnostic characters still preserved in dry as well). Thus, every collection ranks under collection specimens are discovered. C. scalaris a large variety of forms, some- John Eatton LeConte (1824) described times not even presenting these features. Chrysomela scalaris on the basis of color In the volume on of the series and also highlighted two conspicuous elytral Fauna Boreali-Americana, published well markings, namely a moon-shaped humeral over a decade after the description of C. marking and a broad sutural dentate mark- scalaris, its author, Reverend W. Kirby, ing; he also provided an illustration (drawn ignored this species and initiated the confu- by himself; Fig. 1) and mentioned the sion surrounding the taxon (Kirby, 1837). occurrence of the species from New York The description offered by this author for to Florida. These characters were more than Calligrapha philadelphica (Linnaeus, 1758)2 sufficient to distinguish the new species from all other Nearctic Chrysomela Linnaeus, 2 This taxon represents another taxonomic conundrum 1758 known at that time, particularly the which shall be dealt with in a separate study. 2015 TAXONOMIC CLARIFICATION OF CALLIGRAPHA SCALARIS (LECONTE) 3

Figure 3. Lateral, dorsal, and ventral views of ‘‘Chrysomela scalaris LeConte’’ examples in LeConte and Horn Collections (Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts): a–c, Calligrapha ostryae Brown; d–f, C. ignota Brown; g–i, C. floridana Schaeffer; and j–l, C. rowena Knab. Scale bar 5 2.0 mm. fits better the one for C. scalaris or allied 1945 (Fig. 3f). In addition, two varieties were species in highlighting the ‘‘longitudinal mentioned that are currently recognizable as stripe at the suture with three obsolete different species with pale epipleura: one branches’’, a feature clearly missing in C. being perhaps the true C. philadelphica (var. philadelphica. Indeed, he mentioned another B), and one seemingly with the apex of the feature for this species exclusive for one of the arcuate band detached from the subsutural members of the scalaris-group: ‘‘epipleura stripe (perhaps Calligrapha rowena Knab, dark-green’’, only found in C. ignota Brown, 1909; var. C). Even though Kirby’s (1837) 4 BREVIORA No. 541 concept of C. philadelphica did not possibly Boucard; the appearance of this beetle is include the true C. scalaris, Rogers (1857) similar to the C. scalaris-like animals found identified part of the confusion and synon- in northeastern North America and studied ymized the former with LeConte’s taxon, a by Brown (1945), very different from the decision that was retained in all subsequent elm-feeding Calligrapha in southern North catalogues. Rogers (1857) knew Chrysomela America, and thus the Mexican record scalaris LeConte from the Middle States, could be a labeling mistake. Taking a Nebraska, and Lake Superior, and figured a lumper stand to an extreme, and misguided specimen very much fitting the concept for by the confusing taxonomic situation affect- this species used by subsequent authors (but ing Calligrapha, also regarding their ecology see below). Suffrian (1858), in his annotated (e.g., Hagen, 1884), Angell (1885) proposed translation of the previous work, reported that C. scalaris was but a variety or race of this species from many other localities in C. philadelphica by name priority rules. North America such as Louisiana and However, judging from the drawings in this Texas, but also as far south as Mexico article, G. Angell ranked under this name and Costa Rica, obviously misidentifying all North American Calligrapha known at some of the Neotropical species with that time plus several others that would be Calligrapha scalaris. His description of the recognized later. His proposition was criti- sutural marking raises again suspicion of his cized, although interestingly not based on merging under LeConte’s species of several the diagnosis of the existing objective forms later recognized as different species. morphological differences, but on the expe- This author also cited Harris’ (1841) work rience in collecting different-looking animals on New England pest insects to highlight on different plants (Caulfield, 1885). The the association of this species with elm and prevailing taxonomic disorder and the lack linden. In his 13th division of Chrysomela, of knowledge about the importance of in section C (typical North American ecology separating these species jeopardized Calligrapha), Sta˚l (1865) proposed a new biological interpretations affecting them, name, Chrysomela multiguttis, for LeConte’s such as Whitehead’s (1918) very detailed C. scalaris, to avoid homonymy with description of the life cycle of a Chrysomela Doryphora scalaris Olivier, 1807, placed also scalaris LeConte appearing in large num- in the genus Chrysomela Linnaeus under bers on Alnus trees in Nova Scotia. This Sta˚l’s lax taxonomic concept for American author indeed endorsed the idea of this species (Horn, 1884). Immediately feeding upon several trees, not only elms and after, the name fell as synonym and lindens as early proposed by Suffrian (1858). appeared in catalogues under Calligrapha Knab (1909) also associated LeConte’s Calli- scalaris again (e.g., Gemminger and Harold, grapha scalaris with elm and linden, and he 1874; Horn, 1884). Jacoby (1880–1892) was the first to split the C. scalaris (LeConte) reported the species from Mexico and concept to recognize two new taxa: Calligra- offered a comparison with a strikingly pha rhoda Knab, 1909 (Figs. 4d–f) and different southern Nearctic and Neotropical Calligrapha rowena (Figs. 3j–l). This author species, Calligrapha diversa Sta˚l, 1859. The put the emphasis on these species’ food drawing appearing in Biologia Centrali- plants, hazel and dogwood, respectively, as Americana (table 11, fig. 6 in Jacoby, well as several morphological differences. 1888–1892, Suppl.) was based on a speci- Years later, Schaeffer (1928) described men apparently collected in Mexico by M. Calligrapha confluens (Figs. 4a–c), a species 2015 TAXONOMIC CLARIFICATION OF CALLIGRAPHA SCALARIS (LECONTE) 5

Figure 4. Lateral, dorsal, and ventral views of ‘‘Chrysomela scalaris’’ LeConte examples in LeConte and Horn Collection (Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts): a–c, Calligrapha confluens Schaeffer; d–f, C. rhoda Knab; g–i, C. pruni Brown; and j–l, C. dolosa Brown. Scale bar 5 2.0 mm. associated with alder, and clearly divergent C. scalaris in recognizing that the arcuate from C. scalaris. This species was the same band is not bent forming an apical lobe and studied by Whitehead (1918), as easily that most specimens have the suture pale at the deduced from host plant data but also apical declivity of the elytra. The list of type thanks to the superb drawings presented in specimens mentions one individual collected the latter work. The description of C. on alder, the plant where this species lives confluens Schaeffer, 1928 diagnosed it from (Brown, 1945), but the author did not elabo- 6 BREVIORA No. 541

Figure 5. Neotype of Calligrapha scalaris (LeConte) in a, dorsal; b, lateral; and c, ventral views (scale bars 5 2.00 mm), with d, detail of the pronotum (scale bar 5 0.80 mm). rate on this important ecological difference. (1940), who described C. knabi, another The same author described C. scalaris var. Cornus-feeding species, very similar in fact floridana Schaeffer, 1933 (Figs. 3g–i), put- to Knab’s C. rowena, from which it can be ting the emphasis in the comparison with C. recognized by a suite of relatively constant rhoda, perhaps misguided by F. Knap’s characters in both species, including the label attached to the type specimen, ‘‘rhoda basal spot of the arcuate band not com- var. floridana’’, since this species is very pletely fused laterally to the subsutural similar, indistinguishable some may think, stripe in C. knabi Brown, 1940, as well as from the typical C. scalaris. However, the slightly divergent basal ends of paired spots chance that this is a sibling species of C. enclosed by the humeral lunule in this scalaris is possible, as it seemingly feeds on species (generally as U-shaped marking as Cornus (Peck and Thomas, 1998; also, pers. opposed to the more roundish spot in C. obs.), but has also been reported from wax- rowena), or the more numerous and larger myrtle (genus Myrica; Blatchley, 1924). The additional spots on the disc of elytra in next author to recognize a different species Brown’s taxon (compare Fig. 6a with within the group of Calligrapha with broad Fig. 3k). Finally, Brown (1945) made a ‘‘dentate’’ sutural markings was Brown thorough revision of this group, paying 2015 TAXONOMIC CLARIFICATION OF CALLIGRAPHA SCALARIS (LECONTE) 7

Figure 6. Habitus of different species in the ‘‘Calligrapha scalaris’’ group: a, C. knabi Brown (Canada: New Brunswick, Restigouche Co., Kedgwick, 2 June 2012, J. Go´mez-Zurita, H. Viza´n & T. Montelongo leg., IBE-JGZ voucher no. C479); b, C. spiraea (Say) (USA: West Virginia, Greenbrier Co., Tuckahoe Lake, 20 July 2001, J. Go´mez-Zurita leg., IBE-JGZ voucher no. 114); c, living specimen of C. dolosa Brown with pale anterior angles of pronotum (picture by Tom Murray, under Creative Commons License and distributed by BugGuide, http:// bugguide.net); d, lateral view of C. rhoda Knab (USA: New York, Saratoga Co., 7 August 2003, D. J. Funk leg., IBE- JGZ voucher no. 550); and apex of elytra of e, C. virginea Brown (USA: Wisconsin, Brown Co., Bayshore, 24 June 2010, J. Go´mez-Zurita & T. Montelongo, IBE-JGZ voucher no. C134); and f, C. confluens Schaeffer (USA: New York, Essex Co., SW Wilmington, 25 May 2012, J. Go´mez-Zurita, H. Viza´n & T. Montelongo leg., IBE-JGZ voucher no. C505). Scale bars: 2.0 mm. 8 BREVIORA No. 541 particular attention to the biology of the beetle, whereas in the other the circle is , describing seven new species allied to pink. The counterpart in Horn’s collection in- C. scalaris. Interestingly, he had his own idea cludes eight specimens labeled ‘‘Can.’’, three of C. scalaris, not based on the examination of as ‘‘Tex.’’, two as ‘‘N.H.’’, two as ‘‘Neb.’’, one LeConte’s types, and his work based on as ‘‘N.C.’’, one as ‘‘Fla.’’, and one has no northeastern North American beetles possibly locality information. The explicit mention of rendered his elm-feeding C. scalaris not to be several localities in the original description, conspecific with LeConte’s. This was a possi- from New York to Florida, is indicative of the bility contemplated by William Brown, who existence of a syntype series, hopefully includ- hoped that his species ‘‘represents, in part at ed among the specimens at hand, from which least, LeConte’s species’’ (Brown, 1945). thetypespecimenshouldbedesignatedtofix Having had the opportunity to work with the species identity. LeConte’s entomological collection at the The first problem one faces, however, is Museum of Comparative Zoology (MCZ; that these 28 specimens represent as many as Harvard University, Cambridge, Massachu- 9 or 10 different species that have been setts), I tried to clarify in this work some of recognized by subsequent authors on the the taxonomic confusion surrounding the basis of sound morphological differences, as species C. scalaris by justifying the designa- well as by different host plant associations, tion of a type specimen aiming at nomencla- and all of them belong to the C. scalaris tural stability, redescribing the species on complex as identified by Brown (1945). This the basis of this type and providing some situation is revealed to be rather complex by information on how to distinguish the species the fact that LeConte’s ill-defined species from allied taxa in what I present here as the could indeed result in any or several of the ‘‘Calligrapha scalaris’’ group. younger names as synonyms. Not knowing what specimens this author used, it is Neotype designation for Calligrapha possible to make a decision on how to typify scalaris (LeConte) his species preserving the maximum of information that has been accumulating in The conundrum about the identity of C. the literature (except perhaps for C. scalaris scalaris obviously requires the examination of itself), as well as the taxonomic contributions LeConte’s material, something that previous by subsequent authors, preserving the stabil- authors seem to have somehow neglected. The ity of nomenclature. This can be accom- relevant type material is part of the beetle plished by designating as type one of the collection at the MCZ (Harvard Museum of specimens lacking morphological diagnostic Natural History, Cambridge, Massachusetts). characters for other easily recognizable taxa It is currently represented by 28 pinned (not mentioned by LeConte, in any case), specimens distributed between LeConte’s (10 and that does not contradict the currently specimens) and Horn’s collections. The spec- accepted idea of the association of C. scalaris imens used for the original description could with elm, regardless of LeConte’s specimens be in any, as it is generally acknowledged being originally associated with this plant or that these entomologists shared their syntype not, since this information was not available series (P. D. Perkins, MCZ, personal commu- to the author anyway. Proceeding with these nication). None of the specimens in LeConte’s goals in mind, the decision about type collection has locality data; only one specimen designation can be very much simplified has a small blue circle pinned underneath the and the effects on nomenclature reduced. 2015 TAXONOMIC CLARIFICATION OF CALLIGRAPHA SCALARIS (LECONTE) 9

The series of 10 specimens from LeConte’s wings when the is alive. Three of collection shows dark epipleura; two of them Horn’s specimens, one from Canada and (identified with little pink and blue circles) two from New Hampshire, can be ascribed have them brown (Figs. 3a–c), and the other without doubts to C. confluens (Figs. 4a–c), eight black with metallic sheen (Figs. 3d–f). an alder (Alnus) specialist that can be These, together with other less conspicuous recognized by its dark green metallic tint characters, clearly distinguish C. ostryae to head, pronotum, and elytral markings, Brown, 1945 and C. ignota, respectively, from the latter numerous, occupying most of the any other Calligrapha in this group. These elytral surface compared with other species, two species were identified by Brown (1945) showing irregular confluence patterns; sub- as specialists on hop-hornbeam (Ostrya) and sutural stripe separated from sutural stripe birch (Betula), respectively. Five individuals for a relatively long distance basally, and from Canada in Horn’s collection also show the latter usually with a narrow depigment- the black epipleura typical of C. ignota,a ed area along suture on apical declivity of difference that was noted by this author, elytra; spot enclosed by humeral lunule adding a label reading ‘‘epipl. virid.’’ to one of large, ovoid, occupying a large space the individuals. Horn’s specimens also include between lunule, subsutural stripe, and base two species that live associated with dogwood of arcuate band; arcute band at wide obtuse (Cornus), namely Calligrapha floridana (one angle with subsutural stripe, with apical half exemplar, from Florida; Figs. 3g–i) and C. short, generally of irregular profile, perhaps rowena (two individuals, from Canada; confluent with surrounding markings; and Figs. 3j–l). The former, as mentioned above, midlateral spot free from elytral margin. is perhaps the most similar externally The specimen from Nebraska (Figs. 4d–f) among North American species to the elm- no doubt belongs to C. rhoda, distinguished feeding Calligrapha, but its distribution in by a roundish body, dark olivaceous tint to an important area of endemism in North dark areas, and very characteristically by America (Go´mez-Zurita, 2005) and its host the double or transverse spot enclosed by choice seemingly on Cornus allows us to humeral lunule; the shape of its arcuate treat it as a differentiated species, at least band, as two narrow straight arms at wide- until new data become available. Calligra- open obtuse angle, relate it to the form pha rowena is remarkably different from any described as C. rhoda walshiana Knab, 1909. other C. scalaris-like species (except C. C. rhoda feeds exclusively on hazel (Corylus) knabi) in the shape of sutural and subsu- throughout its range. One beetle, the tural stripes, completely fused basally, with smallest in the series and without locality broad blunt lateral expansions below the data, strongly reminds one of the plum tree scutellar area; heavy and basally emargi- (Prunus) specialist, Calligrapha pruni Brown, nated spot enclosed by humeral lunule, 1945, as compared with this species’ para- sometimes confluent with inner margin of types at MCZ (Figs. 4g–i). This species has humeral lunule; arcuate band split in two a rotund body, slender humeral marking, short subtriangular oblique spots, the basal small round spot enclosed by humeral one completely fused to subsutural stripe lunule; rather straight, oblique arms of the and the apical one nearly always free; and arcuate band; relatively large midlateral very reduced additional markings on disc of spot, in most individuals narrowly separated elytra, which are relatively translucent at the from elytral margin; and in the case of the apical two thirds to show the red color of specimen at hand, rather reduced markings 10 BREVIORA No. 541 on elytra. Finally, another specimen in this American Calligrapha since 1824. Yet, also series and labeled ‘‘N. C.’’ is a typical relevant to making a decision, even though Calligrapha dolosa Brown, 1945 (Figs. 4j–l), there is no host information for LeConte’s as compared with Brown’s types, and it has and Horn’s specimens, among the C. sca- been associated with hawthorn (Crataegus). laris-like beetles known to live in Texas, at This specimen does not fit the original least one of them is reportedly feeding on description by LeConte, in not having the elm. There is at least another locally abun- ‘‘suture with a broad greenish blue dentic- dant form that feeds on Cornus (personal ulate line’’, since in this species, and observation; E. G. Riley, TAMUIC, person- particularly in the specimen at hand, the al communication), and it is indistinguish- arcuate band, whose ends produce the able from a morphological point of view ‘‘teeth’’ of the sutural stripe, is clearly from the Ulmus specialist. This could be a detached from this stripe. cryptic species, a host race, or the same For the reasons exposed above, each one species with an exceptionally broad trophic of the specimens mentioned can be discarded range, but at present we cannot ascertain its as potential lectotype for Calligrapha sca- taxonomic status. In any case, by selecting laris. The decision is thus restricted to four one of the Texan specimens in the LeConte specimens, one from Nebraska and three and Horn collections as the Neotype—very from Texas, that still show some differences much consistent with the original description as to be considered two species. Among but also with the posterior usage of the them, the ones from Texas have the greenish name—unless it is possible to demonstrate blue tinge to dark parts mentioned in the that the specimens in the MCZ series actually original description, not so apparent in the belong to a Cornus-feeding species, Calligra- other specimen, which is slightly violaceous. pha scalaris (LeConte) will remain the elm However, the geographic provenance of Calligrapha beetle appearing in scientific these specimens generates an additional literature but also in numerous forestry and problem. Texas (or Nebraska) was not agriculture reports (e.g., Dean, 1946). Be- mentioned in the original description of C. sides, the small, roundish appearance of scalaris. It is not possible to know whether these beetles, as well as their reduced elytral John LeConte had these (or other) specimens markings, particularly the spot enclosed by from Texas at the time he proposed C. humeral lunule, which in two of the individ- scalaris. However, in 1824, the year that the uals is a small round spot (Figs. 5a,b), species was formally described, and until approach the specimen figured by LeConte 1845 these territories were part of Mexico in his original work (Fig. 1a). The drawing and still an exciting frontier of discovery for inexplicably lacks the spot enclosed by American naturalists. Therefore, it is highly lunule, otherwise a very prominent feature unlikely that these specimens were in the in every other C. scalaris-type forms (with entomologist’s hands and that he did not the exception of C. pruni, see Figs. 4g–h and highlight their provenance in his report, compare with Figs. 3g–h, for instance); thus which basically described beetle species it is difficult to understand that the author collected in Georgia (18 of 20). In these missed it if the example used was any of the circumstances, there seems to be no suitable other species. The specimens thus isolated specimen to be designated as lectotype and at to represent LeConte’s species are, again the same time preserving the stability of all with the exception of C. pruni, the smallest subsequent taxonomic additions to North among potential syntypes. LeConte provided 2015 TAXONOMIC CLARIFICATION OF CALLIGRAPHA SCALARIS (LECONTE) 11 data on size, giving half an inch as the size antennae, palpi, legs, internal margin of for this species; however, this information is epipleura, and apical margin of last abdom- not at all reliable, since the largest Calligra- inal ventrite pale orangish brown; pale areas pha known to me is the Mexican C. of elytra creamy yellow. stillatipennis Sta˚l, 1859, nearly twice as big Head large, transverse, deeply inserted into as many North American species, and it does pronotum; surface very finely shagreened, not reach 12 mm. covered by rather dense strong punctures, A sound alternative that would take into except medially on frons before vertex; account distribution information (Monte- punctures near eyes with short anteriorly longo & Go´mez-Zurita, in prep.) and mor- recumbent whitish setae, longer in depressed phological features would be selecting the area above unpunctured supra-antennal specimen from Florida in the MCZ series as calli; frons with fine longitudinal impression lectotype. But this decision would be desta- joining broad V-shaped frontoclypeal su- bilizing. Schaeffer’s C. floridana would auto- ture; area above eyes impressed by furrow matically become a junior synonym of C. prolonged for short distance beyond upper scalaris, which would in turn become asso- eye margin; space behind and below eyes ciated to Cornus, opposing more than coarsely punctured and with relatively long 150 years since the ecological association whitish setae; genae very short. Clypeus with Ulmus was proposed. The elm Calli- transverse, shagreened, strongly punctured grapha would need formal description and a at anterior half, with short, anteriorly new name proposed. Instead, it seems recumbent pale setae; anterior margin at advisable to retain the name associated to sides with longer, medially converging one of the specimens in LeConte’s collection yellowish setae. Eyes relatively small, finely and consistent with its posterior usage. faceted, dorsoventrally elongated, entire. Neotype designation for Calligrapha sca- Labrum transverse, sides curved inward to laris (LeConte). One female, Tex., Horn broadly rounded anterior angles; apex Coll. H6808, Calligrapha scalaris (LeConte) weakly emarginate; transverse row of seti- Neotype des. J. Go´mez-Zurita 2010 (Fig. 5). gerous punctures in front of middle, with Paraneotypes. Two females, Tex., Horn long, medially convergent yellowish setae. Coll. H6808, Calligrapha scalaris (LeConte) Mandibles large, strong, projecting beyond Paraneotype J. Go´mez-Zurita 2010. apical margin of labrum by more than 1.03 its length; sides concave before strong Redescription of Calligrapha preapical curvature; surface densely covered scalaris (LeConte) by large setigerous punctures with long recumbent yellowish setae. Maxillary palpi The Neotype lacks six left apical antenno- slender; second palpomere long, with sub- meres, the terminal segment of right antenna, cylindrical pedicel at basal half, gradually right maxillary palp, left protarsus, and last widening afterward toward apex; third three segments of left mesotarsus. Body oval, palpomere long, base cylindrical, narrower convex, 7.54 mm long, 4.75 mm wide than apex of previous segment, slightly (Figs. 5a–c). Head, most of mandibles, pro- elbowed and strongly widened toward apex; notum, and scutellum black with bluish last segment subtrapezoidal, narrower at metallic tinge; apex of mandibles, elytral base and broadest preapically, before markings, and ventral surfaces dark brown obliquely cut apex. Antennae slender, sub- with slight bluish metallic shine; labrum, clavate beyond fifth antennomere; scape 12 BREVIORA No. 541 elongate oval, thick; pedicel subclavate, straight cut apex; procoxal cavities open twice as long as wide at apex; third posteriorly. Mesanepisterna and mesepimera antennomere slender, twice as long as very finely microgranulose, unpunctured. second; antennomeres 4–6 subequal, as long Metanepisterna shagreened, densely covered as but narrower than pedicel; antennomeres by longitudinally elongated punctures, con- 7–10 short, nearly as long as wide at apex, fluent alongside external margin. Mesoven- strongly widened at base and weakly and trite transverse; space between mesocoxae gradually to apex afterward, densely setose. convex, with fine yellowish setae. Metaven- Pronotum (Fig. 5d) very short, transverse, trite finely leathery, glossy on disc, finely 0.453 shorter medially than wide between impressed longitudinally, with sparse fine posterior angles; basal border at wide obtuse punctures and fine transverse scratches; sides angle, broadly round medially, immargined; finely shagreened, more densely punctured, posterior angles obtuse; sides straight, sub- punctures stronger near anterior angles; parallel, or very weakly convergent at basal surface covered by sparse, very fine, short 2/3, curving toward strongly produced ante- pale yellowish setae. Scutellum 1.143 longer rior angle at apical 1/3; anterior border than broad at base, lancet-shaped, sides concave at angles, nearly straight medially; tapering to blunt pointed apex; surface very sides and apical border finely margined, with finely microgranulose, unpunctured. lateral margin not visible from above right Elytra convex; base as broad as base of behind anterior angles owing to strong pronotum, humeri feebly protruding; sides pronotal convexity at anterolateral declivity; weakly curved, maximum width at middle, surface very finely microreticulated; punctu- apex commonly round; surface very finely ation double on disc, with denser, more or microreticulated, smooth at external interval, less homogeneously distributed minute punc- densely and homogeneously covered by dot- tures and sparse larger punctures, compara- like punctures, dark on paler areas; marginal ble with head punctures; sides of disc marked interval unpunctured, limited internally by slightly ahead of middle by large, deep row of dot-like dark punctures parallel to roundish hollows; punctures at sides stron- elytral margin; punctures larger, deeper ger, deeper, larger than punctures on disc; around and within dark areas. Markings as lateral premarginal area with parallel row of follows (Figs. 5a, b): (i) sutural stripe nar- deep punctures; deep longitudinally elongat- rowly surrounding apical half of scutellum, ed punctures arranged near basal border. narrow, entire, reaching sutural angle; (ii) Hypomera rather flat, uniformly shagreened, basal end of subsutural stripe slightly behind unpunctured, with basal transverse impres- apex of scutellum, weakly divergent and free sions; hypomeral suture obliterated. Proster- from sutural stripe for length of four num short, strongly transverse; anterior punctures, then completely confluent with border concave, margined; disc and anterior it, nearly reaching sutural angle; slightly half at sides microgranulated, unpunctured; widened at apical declivity of elytra before posterior half before procoxae depressed, gradually narrowing to apex; (iii) arcuate with strong, confluent punctures, setose; band consisting of two large spots fused prosternal process convex medially, de- laterally to dark interval confluent with pressed at apex, punctured, with long pale subsutural stripe at middle 1/3 of elytra; yellowish setae mostly at sides and apex; basal spot shaped as elongated inverted tear, process narrow between coxae, as wide as obliquely directed to suture, apical 2/3 antennal scape, gradually widening toward confluent with subsutural stripe; apical spot 2015 TAXONOMIC CLARIFICATION OF CALLIGRAPHA SCALARIS (LECONTE) 13 transverse, bean-shaped, convex apically, gent. Abdominal ventrites leathery on disc, laterally confluent with subsutural stripe; very finely microgranulose at sides, rather (iv) humeral marking consisting of slightly densely punctured by small punctures with curved elongated humeral spot, narrowly short, posteriorly recumbent very fine trans- separated basally from base of elytra, nearly lucent setae; apical segment broadly round, entirely confluent laterally with humeral with narrow premarginal furrow fringed with lunule; lunule elongated, regularly concave erect fine yellowish setae. internally, basally at level with humerus, and apical free end directed to suture and ending What is Brown’s (1945) Calligrapha scalaris? distant but level with basal end of arcuate band; (v) spot enclosed by humeral lunule Fixing the meaning of C. scalaris (Le- small, roundish, placed behind midsection of Conte) to the small, round elm-feeding humeral lunule, 0.53 closer to lunule than to Calligrapha living in central and southern subsutural stripe; (vi) midlateral spot larger U.S.A. legitimates again the question implic- than spot enclosed by lunule, longitudinally it in Brown’s (1945) work. Are the larger, elongated, placed over premarginal line of more slender, more strongly pigmented elm- punctures, occupying five punctures; lateral- feeding Calligrapha in northeastern North ly expanded toward elytral margin, without America conspecific with the C. scalaris as reaching it; (vii) spot of apical declivity as big recognized in the present study? Answering as midlateral spot, roundish, completely this question involves a thorough analysis of fused laterally to preapical enlargement of morphological variation in the species living subsutural stripe; (viii) apical spot round, on Ulmus, and a genetic study would small, equidistant from spot of apical decliv- certainly shed the required light to recognize ity and margin of elytron (it appears as whether these two (or more) forms can be double spot on right elytron of neotype); (ix) considered as the same species, as sister 10 additional small round spots on disc of species, or as independent adaptations to elytra, one on external concavity of arcuate Ulmus as food source. Nonetheless, the degree band, four on lateral declivity evenly spaced of morphological variation observed between between humeral and apical spots, more or specimens reared on elm and named C. less parallel to margin, and five on disc more scalaris by Brown and the C. scalaris based or less evenly spaced in slightly wavy line on Horn’s animals is as high as observed between humeral lunule and spot of apical between different species (as assessed on the declivity. Epipleura smooth, unpunctured. basis of their food choice as well as morpho- Fully developed, reddish wings. logical differences; Brown, 1945). These Femora straight, spindle-shaped, slightly differences may be compatible with treating enlarged medially; surface very finely both forms as different taxa, showing more or shagreened, with sparse small punctures less consistent differences in their elytral and short, adpressed yellowish translucent, markings, e.g., the size and position of the very fine setae. Tibiae straight, gradually spot enclosed by humeral lunule. widening toward apex, as long and similarly The fact that under C. scalaris or more sculptured as femora; apical half with specifically, elm-feeding Calligrapha, there semierect setae longitudinally on edges, are several species that would provide an dense at apex internally. Tarsi slender, explanation for an observation by Robertson shorter than tibiae; first tarsomere shorter (1966) that was difficult to match with our than next two together; claws weakly diver- current knowledge on the biology of these 14 BREVIORA No. 541 beetles. Phylogenetic (Go´mez-Zurita et al., LeConte’s specimens. For instance, Rogers 2006) as well as genetic (Montelongo and (1857) shows a beetle that could represent any Go´mez-Zurita, 2015) studies on these beetles of the C. scalaris-type beetles, but not the reinforce the idea that thelotoky arose southern elm-adapted forms, particularly in several times independently in their evolution that the beetle portrayed is more slender and as a by-product of interspecific hybridiza- it has the spot enclosed by humeral lunule tion. As it occurs with many other animals, large. The same is true for the specimen thelytoky appears in Calligrapha associated figured in Jacoby’s (1880–1892) monograph. with polyploidy (Robertson, 1966). Yet, a Other drawings, particularly the one by E. few species in Calligrapha, including C. Knobel (1895) used by him and subsequent scalaris, were reported as facultatively par- authors to describe the species (e.g., Blatchley, thenogenetic for showing some female-only 1910; Downie and Arnett, 1996), lack suffi- populations or at least those with highly cient detail to make a reliable identification. distorted sex ratios (Brown, 1945; Robert- When ecological information is not avail- son, 1966). Indeed, all other unisexual able, there are at least two species that can be species in the genus are reproductively readily identified thanks to conspicuous isolated from their bisexual counterparts diagnostic character states about the color not only by a different reproductive strategy, of their elytral epipleura. These are C. ignota, but also owing to insurmountable genomic a birch-feeding species that shows blackish imbalances due to the hypothesized polyploid epipleura with metallic sheen (Fig. 3f), and hybrid genomic structure in these species C. ostryae, a hop-hornbeam-feeding species (Montelongo & Go´mez-Zurita, 2015). Thus, with brownish epipleura (Fig. 3c). The next it becomes somewhat of a problem to accept character that is worth looking at is the that this really complex mechanism can be configuration of the midlateral spot (Fig. 2), polymorphic in one species, e.g., C. scalaris.A either broadly confluent with elytral epi- more parsimonious and plausible explanation pleura, or detached from elytral margin (or of these observations is that there are at least narrowly confluent, at most). The specimens two species of Calligrapha associated with identified as C. scalaris have this spot placed Ulmus, probably more, and some of them, medially on elytra, over the midlateral line of including some Canadian and Kansas popu- punctures, expanded toward elytral margin, lations studied by Brown (1945) and Robert- but reaching it narrowly, if at all (Fig. 5b). son (1966), thelytokous. This trait can be important to distinguish the Regardless of differences in their reproduc- elm-feeding Calligrapha from the birch and tive biology, which, if confirmed, would hop-hornbeam species, but also from other strongly support the case of different elm- species that lack the diagnostically dark feeding species, the decision on the taxonomic epipleura, such as C. floridana (Fig. 3g), C. status of Brown’s concept of C. scalaris can be knabi, C. tiliae Brown, 1945, or C. virginea adjourned until new data become available. Brown, 1945, two dogwood and two linden specialists, respectively, otherwise very much ‘‘scalaris’’-like. The midlateral spot is fre- Diagnosis of Calligrapha scalaris (LeConte) quently missing in C. rhoda (Fig. 6d); thus it The drawings for C. scalaris (LeConte) is also a good character to use to diagnose provided by previous authors do not con- this species and differentiate it from C. form to the concept for this species put scalaris and any other species in this group. forward here, on the basis of the study of However, those C. rhoda specimens with 2015 TAXONOMIC CLARIFICATION OF CALLIGRAPHA SCALARIS (LECONTE) 15 midlateral spot can still be easily differenti- small in C. scalaris, C. pruni, and C. rhoda, ated from C. scalaris by their double spot but large, occupying a big proportion of the enclosed by humeral lunule (sometimes both space delimited by humeral lunule, basal end spots are confluent in a U-shaped or of arcuate band, and divergent base of transversal spot), their dark olive-brownish subsutural stripe in the other species, with a tinge to dark parts, as opposed to bluish distance from the spot to any of these features green in C. scalaris, and their stronger body never larger than the diameter of the spot. puncturation, giving a somehow rough ap- With this information, it is possible to pearance to dorsal surfaces (Figs. 4d, e). produce a tentative key to identify the species Another distinctive feature of C. scalaris is in the ‘‘Calligrapha scalaris group’’ on the the well-developed arcuate band, producing basis of practical differences that apply to a the characteristic dentate shape of the sutural large proportion of individuals (at least those marking described by LeConte (1824). The generally matching the phenotype of the presence of this band, with marked basal and respective types), even though not trying to apical lobes, the latter rather big, transverse, reflect the relationship of the species in- sometimes V-shaped (Fig. 5a), is shared by volved. In general terms, it would be possible several species in the group, but readily to recognize the ‘‘C. scalaris group’’ as these distinguishes C. scalaris from C. dolosa and species showing elytra with the subsutural C. rowena (apical lobe detached from sutural stripe confluent with sutural stripe and also stripe; Figs. 3k and 4k), C. rhoda walshiana with one (typically the basal one) or both (apical lobe narrow linear, slightly divergent spots in which the arcuate band is divided from suture, sometimes free; Fig. 4e), C. (except in C. spiraea, where this last feature is confluens (apical lobe straight, short, of free; Fig. 6b). The other group or groups of irregular profile; Fig. 4b), and C. spiraea related North American Calligrapha, tenta- (Say, 1826) (lacks the apical lobe; Fig. 6b). tively proposed here under the name ‘‘Calli- The spot of apical declivity (Fig. 2) on elytra tends to be broadly confluent with subsu- grapha philadelphica’’ group, have these tural stripe in C. scalaris, which should help three features nearly always free (in C. vicina differentiate it again from C. confluens, C. Schaeffer, 1933 and C. suturella Schaeffer, ignota, C. knabi, C. pruni, C. rhoda walshi- 1933 sutural and subsutural stripes are ana, C. rowena, and C. virginea, where this confluent). The key is based in the examina- spot tends to be free. The humeral marking tion of type specimens; thus it is expected to (Fig. 2) in C. scalaris seems smaller, more help identify most individuals. However, compact than in other species in the group, since the proposed key is generally based with the apex of humeral lunule reaching at on the elytral markings and we know that level or only slightly beyond the extreme Calligrapha has a tendency to variation in basal end of the arcuate band (Figs. 5a, b). these markings, it is possible that a certain In other species this marking is more slender, proportion will fail to be keyed out or will reaching for a short distance beyond the actually result in a wrong identification. basal end of the arcuate band, as can be seen in C. confluens, C. ignota, C. pruni, C. tiliae, IDENTIFICATION KEY OF CALLIGRAPHA or C. virginea, and it is heavier, as in C. CHEVROLAT SPECIES IN THE ‘‘CALLIGRAPHA amator Brown, 1945 and most others, except SCALARIS’’ GROUP: perhaps C. pruni. The same is true for the 1a. Midlateral spot absent (Fig. 6d); found on hazel spot enclosed by humeral lunule, relatively (Corylus) ...... Calligrapha rhoda Knab (pars) 16 BREVIORA No. 541

1b. Midlateral spot present (Figs. 3a, d, g, j, 4a, d, g, j, dark body parts; found on elm (Ulmus) and5b) ...... 2 ...... C. scalaris (LeConte) (Fig. 5) 2a. Dark epipleura (Figs. 3c, f) ...... 3 12a. Apical arm of arcuate band generally detached 2b. Pale epipleura (Figs. 3i, l, 4c, f, i, l, and 5c) . . . 4 from subsutural stripe (Figs. 3k, 4k) ...... 13 3a. Black epipleura (Fig. 3f); found on birch (Betula) 12b. Apical arm of arcuate band always confluent with ...... C. ignota Brown (Figs. 3d–f) subsutural stripe (Figs. 4b, 5a, 6a, e) ...... 14 3b. Brown epipleura (Fig. 3c); found on hop-hornbeam 13a. Base of subsutural stripe divergent from sutural (Ostrya) ...... C. ostryae Brown (Figs. 3a–c) stripe; large additional spots on disc of elytra; 4a. Pale anterior angles of pronotum (Fig. 6c); found found on hawthorn (Crataegus) ...... on hawthorn (Crataegus) ...... Calligrapha dolosa Brown (pars) (Figs. 4j–l) ...... Calligrapha dolosa Brown (pars) 13b. Base of subsutural stripe entirely confluent 4b. Dark anterior angles of pronotum (Fig. 5d) . . . . 5 with sutural stripe; small additional spots on 5a. Midlateral spot broadly confluent with margin of disc of elytra; found on dogwood (Cornus) elytron (Fig. 3g) ...... 6 ...... Calligrapha rowena Knab (Figs. 3j–l) 5b. Midlateral spot narrowly confluent or free from 14a. Base of subsutural stripe completely confluent with elytral margin (Figs. 4a, d, g, j, and 5b) ..... 8 sutural stripe; found on dogwood (Cornus) 6a. Spot of apical declivity free from subsutural stripe ...... Calligrapha knabi Brown (Fig. 6a). (Fig. 6e); found on linden (Tilia) ...... 14b. Base of subsutural stripe divergent from sututral ...... C. virginea Brown stripe if for a short distance (Figs. 4b, 5a) . . . . . 15 6b. Spot of apical declivity confluent with subsutural 15a. Suture narrowly pale at apical declivity of elytra stripe ...... 7 (Fig. 6f); found on alder (Alnus) ...... C. confluens Schaeffer (pars) 7a. Humeral marking free from base of elytron (Fig. 5d); found on linden (Tilia) ...... C. tiliae Brown 15b. Suture completely dark along its whole length . . . 16 16a. Apex of arcuate band short, weakly divergent from 7b. Humeral marking broadly connected anteriorly to subsutural stripe, of irregular profile (Fig. 6f), base of elytron (Fig. 3g); found on Cornus usually confluent with surrounding spots; dark green ...... Calligrapha floridana Schaeffer metallic tinge to dark areas; found on alder (Alnus) 8a. Spot enclosed by humeral lunule double, with both ...... C. confluens Schaeffer (pars) (Figs. 4a–c) spots free, fused as a transverse or U-shaped 16b. Apex of arcuate band longer, divergent from marking, but always relatively small (Figs. 4e, 5a, subsutural stripe, V-shaped or more or less neatly 6b,d)...... 9 round; bluish black metallic tinge to dark areas; 8b. Spot enclosed by humeral lunule single, round, oval, found on linden (Tilia)...... C. amator Brown transverse, maybe slightly emarginated anteriorly (Fig. 4h), or if double or U-shaped, then relatively big (Figs. 3k, 6a) ...... 10 9a. Arcuate band confluent with subsutural stripe; On the meaning of Calligrapha multiguttis found on hazel (Corylus) ...... Sta˚l, 1865 ...... Calligrapha rhoda Knab (pars) (Figs. 4d–f) 9b. Arcuate band reduced to two spots free from After several years studying American subsutural stripe; found on ninebark (Physocarpus) Chrysomelinae, the Swedish entomologist ...... C. spiraea (Say) (Fig. 6b) Carl Sta˚l (1865) failed to discover fixed 10a. Spot enclosed by humeral lunule removed from diagnostic differences that unambiguously subsutural stripe by distance over 1.03 width of separated several of the genera dealt with in spot (Figs. 4h, 5a) ...... 11 10b. Spot enclosed by humeral lunule at most as far as his monograph, including Calligrapha. Thus 1.03 its width from subsutural stripe, usually he lumped them under the Linnean genus much less (Figs. 3h, k, 4b, k, 6a, c, e, f) ....12 Chrysomela, resulting in some homonymies 11a. Spot of apical declivity of elytra free or narrowly that he disambiguated. Unfortunately, one connected to subsutural stripe; brownish or bronzy of them affected precisely LeConte’s taxon greenish tinge to head, pronotum, and dark C. scalaris. Sta˚l’s (1865) naming exercise, markings; found on wild plum (Prunus)...... C. pruni Brown (Figs. 4g–i) with the proposition of C. multiguttis to 11b. Spot of apical declivity of elytra generally conflu- represent LeConte’s species, was accompa- ent with subsutural stripe; bluish green tinge to nied by a redescription based on a series of 2015 TAXONOMIC CLARIFICATION OF CALLIGRAPHA SCALARIS (LECONTE) 17 specimens presumably in the beetle collec- apical declivity of elytra, is typical for C. tions of Berlin and Stockholm. It is possible knabi. Fortunately for the stability of to recognize the use of several specimens for current nomenclature, one unlabeled spec- this redescription based on the mention of imen and the one closing the series, with at least two depositories, and it becomes the label ‘‘scalaris Rog. Suff.*, Texas, clear that more than one specimen was used Friedr.’’, have very reduced markings, according, for instance, to the description typical of C. scalaris as interpreted in this of the spot of apical declivity close to work, particularly by the spot enclosed by subsutural stripe or sometimes (interdum) humeral lunule small and conspicuously confluent with it. Knowing the circum- closer to humeral lunule than to subsutural stances around the name C. scalaris (Le- stripe, as well as free midlateral spot. Conte), it seems important to try to However, these specimens show a free spot understand Sta˚l’s concept of this species, of apical declivity, approaching C. pruni and therefore the meaning of C. multiguttis. instead, and in any case cautioning about I could not localize the specimen(s) at the the use of this character as diagnostic of Naturhistoriska riksmuseet (Stockholm, these taxa. With this information at hand it Sweden), which hosts Sta˚l’s collection. They is possible to split the synonymy for the could have been arranged among other C. different taxa involved: C. scalaris (Le- scalaris by Sta˚l himself, curator of this part Conte) (5 Chrysomela multiguttis Sta˚l, of the collection. However, the original 1865 [pars], Mon. Chrys. Am., iii, p. 261), series has been preserved and identified at C. ignota Brown (5 Chrysomela multiguttis the Museum fu¨r Naturkunde (Berlin, Ger- Sta˚l, 1865 [pars] nov. syn.,Mon.Chrys. many), slightly complicating the picture of Am., iii, p. 261), C. knabi Brown (5 what represents the name C. multiguttis and Chrysomela multiguttis Sta˚l, 1865 [pars] how it relates to C. scalaris. Indeed, the nov. syn., Mon. Chrys. Am., iii, p. 261), examination of this series of six specimens and C. tiliae Brown (5 Chrysomela multi- (Nr. 29806 in the MfN Collection registry) guttis Sta˚l, 1865 [pars] nov. syn., Mon. reveals that it includes at least five species, Chrys. Am., iii, p. 261). all of them in what could be considered the ‘‘scalaris-complex’’, similarly as it happens ACKNOWLEDGMENTS with LeConte’s and Horn’s collections. The series opens with one specimen labeled as My deepest thanks go to Dr. Brian D. ‘‘29806, scalaris Rog. Suffr.*, Amer. Sept.’’, Farrell and Dr. Philip D. Perkins (MCZ, which fits the description and appearance Harvard) for their support and for so kindly of C. floridana, with midlateral spot broad- hosting me during my stay at MCZ. Phil ly confluent with elytral margin, as well as shared his knowledge about the singularity spots of apical declivity fused with sub- of LeConte’s collection, as well as useful sutural stripe. An unlabeled specimen with information and valuable opinions that similar characteristics but with the humeral helped me form the idea for this work. marking free from elytral margin basally Thanks also to Tinguaro Montelongo (Insti- could be referred to C. tiliae.Another tute of Evolutionary Biology, Barcelona) for unlabeled specimen has conspicuous black the time spent together on entomological epipleura; thus it belongs in C. ignota.Yet campaigns in North America collecting C. another unlabeled specimen, smaller, with scalaris and the other species and for large midlateral spot and free spots on discussion on species meaning and limits in 18 BREVIORA No. 541 this wonderful evolutionary lineage of bee- GO´ MEZ-ZURITA, J. 2005. New distribution records and tles. Dr. Piotr Nakrecki (MCZ) spared his biogeography of Calligrapha species (leaf beetles), in North America (Coleoptera: Chrysomelidae, Chry- digital imaging facility to illustrate this work. somelinae). Canadian Field-Naturalist 119(1): 88–100. The study of the meaning of C. multiguttis GO´ MEZ-ZURITA, J., D. J. FUNK, AND A. P. VOGLER. 2006. (Sta˚l) was possible thanks to the FP6 EU The evolution of unisexuality in Calligrapha leaf Synthesys Programme projects SE-TAF-1893 beetles: Molecular and ecological insight on multi- (Naturhistoriska riksmuseet, Stockholm) and ple origins via interspecific hybridization. Evolution 60: 328–347. DE-TAF-4348 (Museum fu¨r Naturkunde, GO´ MEZ-ZURITA, J., A. P. VOGLER, AND D. J. FUNK. 2004. Berlin). Brian Farrell and two anonymous Diagnosing an overlooked North American taxon: referees contributed with their comments and Biological observations and mitochondrial insights suggestions to improve some important on Calligrapha suturella Schaeffer, new status weaknesses in my original submission, help- (Coleoptera, Chrysomelidae). Annals of the Ento- mological Society of America 97: 28–36. ing me to present a stronger case. This study HAGEN, H. A. 1884. 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Fauna Boreali- of the Rhynchophora) Known to Occur in Indiana. Americana; or the Zoology of the Northern Parts of Indianapolis, Nature Publishing Co. British America Norwich, U.K., Josiah Fletcher. BLATCHLEY, W. S. 1924. The Chrysomelidae of Florida. KNAB, F. 1909. Some species of Calligrapha (Coleoptera, Florida Entomologist 8(1): 1–7. Chrysomelidae). Proceedings of the Entomological BROWN, W. J. 1940. Some new species of Cantharidae Society of Washington 11: 83–87. and Chrysomelidae. Canadian Entomologist 72(8): KNOBEL, E. 1895. Beetles of New England and Their Kind. 161–166. A Guide to Know Them Readily Boston, Bradlee BROWN, W. J. 1945. Food-plants and distribution of the Whidden. 48 pp. species of Calligrapha in Canada, with descriptions LECONTE, J. E. 1824. Descriptions of some new species of new species (Coleoptera, Chrysomelidae). Cana- of North American insects. Annals of the Lyceum of dian Entomologist 77(7): 117–133. Natural History of New York 1: 169–173. CAULFIELD, F. B. 1885. Notes on Chrysomela scalaris, LINNAEUS, C. 1758. Systema Naturae per Regna Tria Lec. Canadian Entomologist 17(11): 230. Naturae, secundum Classes, Ordines, Genera, Spe- DEAN, G. A. 1946. The elm Calligrapha (Calligrapha cies, cum Characteribus, Differentiis, Synonymis, scalaris Lec.). Agricultural Experiment Station, Kansas Locis Tomus I. Holmiae, Laurentii Salvii. 823 pp. State College, Manhattan, Kansas, Circular 234: 1–7. MONTELONGO, T., AND J. GO´ MEZ-ZURITA. 2013. Mor- DOWNIE,N.M.,AND R. H. ARNETT. 1996. The Beetles of phological and molecular characterization of a new Northeastern North America. Volume II: Polyphaga: Nearctic species of Calligrapha Chevrolat, 1836 Series Bostrichiformia through Curculionoidea Gaines- (Coleoptera: Chrysomelidae, Chrysomelinae) from ville, Florida, Sandhill Crane Press. pp. i–x, 891– Central Mexico. Proceedings of the Entomological 1721. Society of Washington 115: 369–391. GEMMINGER,M.,AND E. vON HAROLD. 1874. Catalogus MONTELONGO, T., AND J. GO´ MEZ-ZURITA. 2014. Multi- Coleopterorum hucusque descriptorum synonymi- locus molecular systematics and evolution in time cus et systematicus. Tom. XI. Chrysomelidae (Pars and space of Calligrapha (Coleoptera: Chrysomeli- I.). Monachii: Sumptu G. Beck, pp. 3233–3478. dae, Chrysomelinae). Zoologica Scripta 43: 605–628. 2015 TAXONOMIC CLARIFICATION OF CALLIGRAPHA SCALARIS (LECONTE) 19

MONTELONGO, T., AND J. GO´ MEZ-ZURITA. 2015. Non- SCHAEFFER, C. 1928. The North American species of random patterns of genetic admixture expose the Hydrothassa with notes on other Chrysomelidae and a complex historical hybrid origin of unisexual leaf description of new species and a variety (Col.). Journal beetle species in the genus Calligrapha. American of the New York Entomological Society 36: 287–291. Naturalist 185: (in press). SCHAEFFER, C. 1933. Short studies in the Chrysomelidae OLIVIER, A. G. 1807. Entomologie, ou Histoire Naturelle (Coleoptera). Journal of the New York Entomolog- des Insectes. Cole´opte`res. Tome Cinquie´me Paris, ical Society 41: 457–480. Chez Desray. 612 pp. STA˚ L, C. 1862–1865. Monographie des Chrysome´lides de PECK, S., AND M. THOMAS. 1998. A Distributional l’Ame´rique Uppsala, Sweden, C. A. Leffler. 365 pp. Checklist of the Beetles (Coleoptera) of Florida. SUFFRIAN, C. W. L. E. 1858. Uebersicht der in den of Florida and Neighboring Land Areas. Vereins, Staaten von Nord-Amerika einheimischen Volume 16 Gainesville, Florida Department of Chrysomelen. Entomologische Zeitung herausgege- Agriculture and Consumer Services. 180 pp. ben von dem entomologischen Vereine zu Stettin ROBERTSON, J. G. 1966. The chromosomes of bisexual and 19(7–9): 237–278. parthenogenetic species of Calligrapha (Coleoptera: WHITEHEAD, W. E. 1918. Notes on the life history and Chrysomelidae) with notes on sex ratio, abundance immature stages of three common chrysomelids. and egg number. Canadian Journal of Genetics and Proceedings of the Entomological Society of Nova Cytology 8: 695–732. Scotia 4: 38–50. ROGERS, W. F. 1857. Synopsis of species of Chrysomela WILCOX, J. A. 1972. A Review of the North American and allied genera inhabiting the United States. Chrysomeline Leaf Beetles (Coleoptera: Chrysome- Proceedings of the Academy of Natural Sciences of lidae). New York Museum and Science Service Philadelphia 8[1856] 29–39. Bulletin No. 421. 37 pp.