Arctodium vulpinum ( Erichson, 1835 )

Hawkins, Shauna Joy, 2006, A revision of the Chilean tribe Lichniini Burmeister, 1844 (Coleoptera: Scarabaeidae: Melolonthinae), Zootaxa 1266 (1), pp. 1-63 : 51-60

Arctodium vulpinum ( Erichson, 1835)

( Figs. 2 , 6a , 7a , 8c , 12 )

Arctodium villosum Dejean 1833: 167 (nomen nudum).

Arctodium vulpinum ( Erichson 1835: 269) (valid name).

Cratoscelis vulpina Erichson 1835: 269 (original combination).

Cratoscelis aterrima Blanchard 1850: 53 . New synonymy.

Cratoscelis gayana Blanchard 1850: 53 . New synonymy.

Cratoscelis villosa Blanchard 1850: 53 . New synonymy.

Cratoscelis striolata Redtenbacher 1868: 61 . New synonymy.

Cratoscelis obscura Germain 1911: 68 (nomen nudum).

Cratoscelis margine­costata Germain 1911: 68 (nomen nudum).

Catalog

Arctodium villosum: Dejean 1833: 167 (first usage, without description, catalog); Dejean 1836: 186 (catalog); Burmeister 1844: 10 (catalog listing as synonym of Cratoscelis vulpina Erichson ); Lacordaire 1856: 162 (checklist).

Arctodium vulpinum: Chapin 1938: 81 (transferred to Arctodium Burmeister from Cratoscelis Erichson ); Hawkins 2004 (online catalog).

Cratoscelis vulpina: Erichson 1835: 269 , Pl.3, Fig.5 ; Laporte 1840: 155; Burmeister 1844: 10 (redescription); Curtis 1845: 455 (catalog, note on mouthparts); Blanchard 1850: 52 (catalog); Solier 1851: 119–122, Plate 17, Fig. 10 (redescription and illustrations); Lacordaire 1856: 162 (checklist); Philippi 1887: 685 (catalog); Redtenbacher 1868: 61 (catalog); Harold 1869: 1094 (catalog); Germain 1911: 68 (catalog); Arrow 1912: 6 (catalog); Chapin 1938: 80 (designated as type species of Arctodium ); Blackwelder 1944: 216 (checklist); Machatschtke 1959: 530, 532, 541 (illustrations); Hawkins 2004 (online catalog).

Cratoscelis aterrima: Blanchard 1850: 53 (original description); Solier 1851: 122 (redescription and diagnoses); Lacordaire 1856: 162 (checklist); Philippi 1887: 685 (catalog); Redtenbacher 1868: 61 (catalog); Harold 1869: 1094 (catalog); Germain 1911: 68 (catalog); Arrow 1912: 6 (catalog); Blackwelder 1944: 216 (checklist); Machatschtke 1959: 530, 532, 533 (illustrations), 539 (distribution map); Hawkins 2004 (online catalog).

Cratoscelis gayana Blanchard 1850: 53 (original description); Solier 1851: 121,122 (redescription and diagnosis); Lacordaire 1856: 162 (checklist); Philippi & Philippi 1864: 326 (usage, comparison with Cratoscelis canicapilla ); Harold 1869: 1094 (catalog); Philippi 1887: 685 (catalog); Germain 1911: 68 (catalog); Arrow 1912: 6 (catalog); Blackwelder 1944: 216 (checklist); Machatschtke 1959: 530, 532, 533 (illustrations), 539 (distribution map); Hawkins 2004 (online catalog).

Cratoscelis villosa: Blanchard 1850: 53 (original description); Solier 1851: 121 (redescription and diagnoses); Philippi & Phillip 1864: 326 (usage, comparison with Cratoscelis canicipilla ); Lacordaire 1856: 162 (checklist); Harold 1869: 1094 (catalog); Philippi 1887: 685 (catalog); Germain 1911: 68 (catalog); Arrow 1912: 6 (catalog); Blackwelder 1944: 216 (checklist); Machatschtke 1959: 530, 532, 533 (illustrations), 539 (distribution map); Hawkins 2004 (online catalog).

Cratoscelis striolata: Redtenbacher 1868: 61 (original description); Philippi 1887: 685 (catalog); Harold 1869: 1094 (catalog); Germain 1911: 68 (catalog); Arrow 1912: 6 (catalog); Blackwelder 1944: 216 (checklist); Machatschtke 1959: 530, 532, 533 (illustrations), 539 (distribution map); Hawkins 2004 (online catalog).

Cratoscelis obscura: Germain 1911: 68 (first usage, without description, catalog); Hawkins 2004 (online catalog).

Cratoscelis margine­costata: Germain 1911: 68 (first usage, without description, catalog); Hawkins 2004 (online catalog).

Type specimens

Cratoscelis vulpina , lectotype male at ZMHB labeled: a) “ vulpina Er. ” (handwritten), b) “ vulpina / Erichs./ Chili” (large, green, square label, handwritten), c) “ Southern Neotropical Scarabs / database #SH75981045/ Arctodium vulpinum Erichson ♂ / DET: S.J. Hawkins 2005” (typeset, ♂ hand­drawn), d) “ Cratoscelis vulpina / Erichson/ LECTOTYPE ♂ / S.J. Hawkins ” (red label, typeset). Lectotype here designated. Erichson (1835) did not indicate how many specimens were in the type series. The location and existence of paralectotypes is unknown. Type locality: Chile .

Cratoscelis aterrima Blanchard , lectotype female at MNHM labeled: a) “7/ 49” (round green label, handwritten on underside), b) “ C. aterrima ./ Cat. mus./ Chili/ M. Gay. ” (large green label, handwritten), c) “ Valparaiso / 72,” d) “Southern Neotropical Scarabs/ database #SH75982035/ Arctodium vulpinum Erichson ♀ / DET: S.J. Hawkins 2005” (typeset, ♀ hand­drawn), d) “ Cratoscelis aterrima / Blanchard/ LECTOTYPE ♀ / S.J. Hawkins ” (red label, typeset). Lectotype here designated. One female paralectotype at MNHN labeled: a) “4196/ 34” (round green label, handwritten on underside), b) “Southern Neotropical Scarabs/ database #SH7592036/ Arctodium vulpinum Erichson ♀ / DET: S.J. Hawkins 2005” (typeset, ♀ hand­drawn), c) “ Cratoscelis aterrima / Blanchard/ PARALECTOTYPE ♀ / S.J. Hawkins (yellow label, typeset). One female paralectotype at MNHN labeled: a) “15/ 45” (round green label, handwritten on underside), “Southern Neotropical Scarabs/ database #SH7592037/ Arctodium vulpinum Erichson ♀ / DET: S.J. Hawkins ” (typeset, ♀ hand­drawn), c) “ Cratoscelis aterrima / Blanchard/ PARALECTOTYPE ♀ / S.J. Hawkins (yellow label, typeset). One female paralectotype at MNHN labeled: a) “15/ 45” (round green label, handwritten on underside), c) “ Cratoscelis ????” (sic) (handwriting not legible), d) “Southern Neotropical Scarabs/ database #SH7592038/ Arctodium vulpinum Blanchard ♀ / DET: S.J. Hawkins 2005” (typeset, ♀ hand­drawn), e) “ Cratoscelis aterrima / Blanchard/ PARALECTOTYPE ♀ / S.J. Hawkins (yellow label, typeset). One male paralectotype at MNHN labeled: a) “4/ 49” (round green label, handwritten on underside), b) “Southern Neotropical Scarabs/ database #SH75982039/ Arctodium vulpinum Erichson ♂ / DET: S.J. Hawkins 2005” (typeset, ♂ hand­drawn), c) “ Cratoscelis aterrima / Blanchard / PARALECTOTYPE ♂ / S.J. Hawkins (yellow label, typeset). The specimens in the type series were obtained from Claude Gay. Blanchard (1850) did not indicate how many specimens were in the type series. The location and existence of other paralectotypes is unknown. See remarks section for an explanation of synonymy. Type locality: Valparaíso, Region V (Valparaíso), Chile.

Cratoscelis gayana Blanchard , lectotype female at MNHM labeled: a) “15/ 45” (round green label, handwritten on underside), b) “ Coquimbo ” (handwritten) c) “ C. gayana / Cat. Mus./ Chili/ M. Gay ” (large green label, handwritten), d) “Southern Neotropical Scarabs/ database #SH75982032/ Arctodium vulpinum Erichson ♀ / DET: S.J. Hawkins 2005” (typeset, ♀ hand­drawn), e) “ Cratoscelis gayana / Blanchard/ LECTOTYPE ♀ / S.J. Hawkins ” (red label, typeset). Lectotype here designated. Two female paralectotypes at MNHN labeled: a) “15/ 45” and “670/ 97” (round green label, handwritten on underside), b) “Southern Neotropical Scarabs/ database #SH75982033/ Arctodium vulpinum Erichson ♀ / DET: S.J. Hawkins 2005” and “Southern Neotropical Scarabs/ database #SH75982034/ Arctodium vulpinum Erichson ♀ / DET: S.J. Hawkins 2005” (typeset, ♀ hand­drawn), c) “ Cratoscelis gayana / Blanchard / PARALECTOTYPE ♀ / S.J. Hawkins (yellow label, typeset). Blanchard (1850) did not indicate how many specimens there were in the type series. The location and existence of other paralectotypes is unknown. See remarks section for an explanation of synonymy. Type locality: Region IV (Coquimbo), Chile.

Cratoscelis villosa Blanchard , lectotype female at MNHM labeled: a) “3065/ 40” (round green label, handwritten on underside, b) “ C. villosa ./Dej./ Chili./ M. Gay. ” (large green label, handwritten), c) “Southern Neotropical Scarabs/ database #SH75982031/ Arctodium vulpinum (Erichson) ♀ / DET: S.J. Hawkins 2005” (typeset, ♀ hand­drawn), d) “ Arctodium villosum / Blanchard/ LECTOTYPE ♀ / S.J. Hawkins ” (red label, typeset). Lectotype here designated. Blanchard (1850) stated that he based his concept of C. villosa on females but did not indicate how many specimens were in the type series. The location and existence of other paralectotypes is unknown. See remarks section for an explanation of synonymy. Type locality: Chile.

Cratoscelis striolata Redtenbacher , lectotype male at NMWH labeled: a) “ Zelebor / Novara ” (typset), b) “Z” (typeset), c) “ Type ” (handwritten), d) “TYPUS” (red, rectangular label, typeset), e) “ Striolata / Chili. Rdt ” (rectangular label with octaganol, black border, handwritten), f) “ Southern Neotropical Scarabs / database # AS 2603128 / Arctodium vulpinum (Erichson) ♀ / DET: A.B.T. Smith 2006” (typeset), g) “ Cratoscelis striolata / Redtenbacher/ Shauna Hawkins/ Lectotype / ♀ ” (red label with black border, handwritten). Lectotype here designated. Paralectotype male at NMWH labeled: a) “Zelebor/ Novara” (typset), b) “Z” (typeset), c) “TYPUS” (red, rectangular label, typeset), d) “ Cratoscelis / striolata / Type. Redtb.” (handwritten), e) “ Southern Neotropical Scarabs / database #AS2603129/ Arctodium vulpinum (Erichson) ♀ / DET: A.B.T. Smith 2006” (typeset), f) “ Cratoscelis striolata / Redtenbacher/ Shauna Hawkins/ Paralectoype ♀ ” (handwritten). The location and existence of other paralectotypes is unknown. See remarks section for an explanation of synonymy. Type locality: Chile.

Description

Male. Length 8.0–12.0 mm; width 5.0– 7.6 mm. Color: Head, pronotum, scutellum , and pygidium black; venter and legs reddish brown to black; elytra dull brown to chestnut brown. Head: Surface punctate; punctures small, dense, setose; setae moderately long, moderately dense to dense, golden yellow to golden brown. Labrum with surface punctate; punctures small, dense, setose; setae short to moderately long, half as long as clypeal setae, dense, golden brown to black. Mandibular external surface with short to moderately long, dense, black setae. Maxillary palpi with short, black, apical setae on all segments except last. Labial margins with moderately long, dense, black setae. Labial palpi with short, black setae at apex of all segments except last. Eye canthus setose; setae long, dense, black. Antenna with pedicel setose, elongate, segment 2 globular, 3–6 subequal in length, 7–9 lamellate; club subequal in length to segments 3–6 ( Fig. 4b ). Pronotum: Surface punctate; punctures small, dense, with long, dense, golden yellow to golden brown setae. Anterior margin setose; setae short, dense, golden brown. Lateral margins setose; setae long, dense, golden yellow to golden brown. Posterior margin setose; setae short to moderately long, dense, golden yellow to golden brown. Scutellum: Surface setose; setae moderately long, moderately dense, golden brown. Elytron: Form moderately wide, less than half as wide as long. Surface flat, punctate; punctures small, moderately dense medially, dense laterally, setose; setae moderately long, golden yellow to golden brown. Propygidium: Surface exposed, covered with long, dense, white setae; setae overlaying basal 1/5 of pygidium (covered by elytra in some specimens), apices of setae forming straight row. Pygidium: Surface slightly concave apically, setose; setae long, dense, golden brown, dark brown, or black. Venter: Thorax, mes­ and metepisternum punctate; punctures small, dense, with golden brown setae. Abdominal segments densely setose on lateral margin, sparsely setose medially; setae long, golden brown. Legs: Surface setose, setae long, dense, golden yellow to golden brown. Protibia slender, with 3 outer teeth and 2 inner teeth. Protibial spur small, 3/4 length of first tarsal segment. Metafemur broadly expanded. Metatibial apex with 2 spurs, 1 spine; spine between 2 spurs, longer than spurs, oblique at apex ( Fig. 8c ). Parameres: Fig. 9c .

Female. Females differ from males in the following respects: length 8.5–11.7 mm; width 4.5–6.2 mm. Color: Elytra, venter, legs, and setae as in males or elytra, venter, legs, and setae black in black female color morph. Elytron: Surface convex, not flat or depressed. Legs: Metatibial apex with 1 spine; spine shorter than adjacent spurs, broadly attenuate at apex ( Fig. 8d ).

Diagnosis

Arctodium vulpinum is the most widespread and common species in the tribe Lichniini . It has a broad range of variation in morphological characters that make it a difficult species to diagnose. Arctodium vulpinum is, on average, the largest species of Lichniini , with males 8.0–12.0 mm long and females 8.5–11.7 mm long. Because there is some overlap in size (smaller specimens of A. vulpinum are as small as the largest specimens of A. discolor or A. planum ), the best way to distinguish small specimens is by ruling out characters defining other species: A. vulpinum does not have a pale fringe of setae along the eye canthus as in A. planum , it does not have black setae on the legs and wide elytra giving it a square­like body form as in A. discolor , or it is not less than 6.5 mm long as in A. mahdii .

Remarks

In Blanchard’s (1850) description of Cratoscelis aterrima , he differentiated the species from others in the genus on the basis of the black color on all parts of the body. However, this species is based on a black color morph found in females of A. vulpinum . Aside from its black color, C. aterrima shares all other characters with A. vulpinum . I observed the more common brown morph males mating with black female morphs on flowers in Cajon del Maipo, Chile. In collections, series of A. vulpinum contain approximately one female black morph out of every eight female specimens collected.

Although black color morphs have not been named as a subspecies in any publication, many collections have specimens labeled as C. vulpina aterrima . This is in keeping with my hypothesis that the black specimens should be classified as A. vulpinum , but I do not believe that black color morphs constitute a subspecies. Mayr (1963) defines a subspecies as “an aggregate of local populations of a species inhabiting a geographic subdivision of the range of the species, taxonomically separated from other populations of the species.” Based on this definition, C. aterrima is not a subspecies because it is not an isolated subdivision of A. vulpinum but is found throughout the populations as a phenotypic variant.

Blanchard (1850) described Cratoscelis gayana based on the pattern of punctures on the elytra. According to Blanchard, the punctures of C. gayana form ordered, straight rows, whereas the punctures of other species of Cratoscelis are random. However, all species of Lichniini have at least one row of punctures that form a straight row at the inner margin of the elytra. Laterad from the inner margin of the elytra, the punctures become gradually more random. The extent to which the punctures become more random is variable from individual to individual. I have studied Blanchard’s type series consisting of three females deposited in the MNHN and have noted that the rows Blanchard is referring to are not as ordered into straight lines as his description indicates, and they are not more so than in specimens of A. vulpinum . Furthermore, in specimens where the setae are rubbed off (from collection, rough handling, or due to the age of the specimen), as is the case with Blanchard’s type series, punctures that seem to form rows seem more pronounced because the pattern of the punctures is more visible. If Blanchard was comparing specimens where the elytral setae were rubbed off with those where the setae were still intact, he might have erroneously divided the specimens into two species. Other characters Blanchard used to define C. gayana , such as body length, pattern and color of setae, and body shape, are consistent with characters for A. vulpinum . Therefore I conclude that C. gayana is conspecific with A. vulpinum .

Blanchard (1850) created the unnecessary name Cratoscelis villosa when trying to validate Dejean’s species name Arctodium villosum , which was published without a description (see the Taxonomic History section). The name A. villosum Dejean was already synonymized with C. vulpina Erichson by Burmeister (1844). Because Blanchard (1850) was the first to publish a description in connection with the name Cratoscelis villosa , it is a valid name that I here synonymize with A. vulpinum . I also confirmed this by studying Blanchard’s single female type specimen deposited at MNHN.

In Redtenbacher’s (1868) description of Cratoscelis striolatum , all characters described are consistent with those of A. vulpinum in terms of length, width, color, shape, and setae. Redtenbacher claims to have studied other species of Cratoscelis and differentiates C. striolatum from other species in the genus based only on punctures, especially of the pronotum, which he stated are dense in C. striolatum , with “gaps between the punctures not larger than the punctures themselves” (my translation). However, this is also true of the punctures of the pronotum of specimens of other species and a fine, linelike pattern can be observed to a varying degree in all species based on how clean the pronotum is and the quality of the light source. I conclude that C. striolata is conspecific with A. vulpinum .

While Germain’s (1911) two names, Cratoscelis obscura and Cratoscelis marginecostata , are nomena nuda (see Taxonomic History section), I mention them here because

these names have been perpetuated in collections. Germain published these names without description in his catalog of the Coleoptera of the Museum of Chile. I have studied the Germain specimens at MNNC, and I conclude that he used the name C. marginecostata for brown morphs of A. vulpinum and C. obscura for black morphs of A. vulpinum . Germain did not describe any new species, and the names remain nomina nuda. Germain’s specimens correspond to numbers in his catalog: 1787 is on the label of the specimen he examined and on which he based the name C. margine­costata , and 1788 is on the C. obscura label.

Distribution

Widespread in central Chile from Region IV (Coquimbo) to Region IX (La Araucania) ( Fig. 12 ) .

Locality data

943 specimens were examined from BMNH, CASC, CMNC, CNCI, FMNH, HAHC, JEBC, JMEC, KSUC, LACM, LEMQ, MCZC, MNHN, MNNC, UCCC, UMRM, UNSM, USNM, VMDC, ZMHB, ZSMC .

CHILE. REGION IV (COQUIMBO) (77): Hurtado (1), Illapel (69), Socos (7). REGION V (VALPARAÍSO) (72): Aconcagua (7), Alfalfares (10), Algarrobo (5), Canelillos (13), Colliguay (2), El Guindal (3), El Quisco (1), El Salto (2), La Campana (2), Las Peñas (1), Limache (6), Llo­Lleo (2), Punta Horcón (1), Rodelillo (1), Vina del Mar (2), Zapallar (3), no data (11). REGION METROPOLITANA DE SANTIAGO (467): Alhue (1), Bella Vista (1), Cajon del Maipo (42), Cerro Manquehue (2), Chacabuco (12), Conchali (1), Cordillera El Tollo (3), Cuesta Barriga (5), El Canelo (40), El Clarillo (1), El Ingenio (1), El Manzano (25), El Noviciado (2), El Peumo (27), El Portezuelo (1), El Principal (4), El Tabo (3), Farellones (2), Florida (2), La Africana (15), La Obra (19), Laguna Carén (6), Lampa (6), Lo Marin (1), Macul (3), Maipu (30), Peñalolén (3), Pudahuel (63), Quebrada de San Ramon (29), Renca (13), Rio Clarillo (3), Rio Colorado (2), Rio Maipo (10), San Bernardo (16), San José de Maipo (5), Santiago (31), Talagante (4), Tiltil (4), Valle Ramon (33), Vizcachas (11). REGION VI (O’HIGGINS) (10): Las Nieves (1), Parral (5), Rancagua (2), Tanumé (1), Termas de Cauquenes (1). REGION VII (MAULE) (24): Constitucion (6), Curicó (1), Fundo Tregualemu (1), La Jaula (1), Pelluhue (1), Peunte Malcho (3), Rio de los Cipreses (5), San Clemente (3), Talca (3). REGION VIII (BIOBÍO) (38): Chillan (1), Cobquecura (1), Concepción (21), Contulmo (7), El Salto del Laja (1), Mulchen (3), Nahuelbuta (1), Quilaco (1), Talcahuano (1), Temuco (1). REGION IX (LA ARAUCANIA) (4): Angol (1), Horcones (2), Icalma (1), Victoria (1). NO DATA (250).

Temporal data January (98), February (22), October (140), November (268), December (243).

Figures

FIGURE 2. Arctodium vulpinum (Erichson) male.

FIGURES 4a–c. Dorsal view of antenna showing length of club. a) Lichnia limbata male; b) L. limbata female; c) Arctodium mahdii male.

FIGURES 5a–b. Dorsal view of maxilla showing length of galea. a) Arctodium discolor; b) Lichnia limbata.

FIGURES 6a–b. Dorsal view of labrum and mandibles showing form of the mandibles (a) dentate, Arctodium vulpinum or (b) rounded, Lichnia limbata.

FIGURES 7a–b. Lateral view of beetles showing length of galea. a) Arctodium vulpinum; b) Lichnia limbata.

FIGURES 8a–d. Metatibial apex showing apical spine (arrow) and adjacent spur. a) Arctodium planum male; b) A. mahdii male; c) A. vulpinum male; d) A. vulpinum female.

FIGURES 9a–c. Caudal (i) and lateral (ii) views of parameres. a) Lichnia gallardoi; b) L. limbata; c) Arctodium discolor.

FIGURE 10. Map of central Chile showing distribution of Lichnia gallardoi and L. limbata.

FIGURE 12. Map of central Chile showing distribution of Arctodium vulpinum and A. mahdii.