Cylindera (Cylindera) morio ( Klug, 1834 )

Moravec, Jiří & Šafránek, Ondřej, 2025, Taxonomic revision of the current concept of Cylindera morio and allied taxa (Coleoptera: Cicindelidae), Acta Entomologica Musei Nationalis Pragae 65 (1), pp. 105-147 : 110-114

Cylindera (Cylindera) morio ( Klug, 1834)

( Figs 1–2 , 11–72 )

Cicindela morio Klug, 1834: 16 . Cicindela egena Chaudoir, 1854: 12 (synonymy by FLEUTIAUX 1892: 68). Cicindela morio var. egena : CHAUDOIR (1865: 40). Cicindosa morio : SCHILDER (1953: 561). Cylindera (Cylindera) morio : RIVALIER (1954: 265). Cicindela (Cylindera) morio morio : SUMLIN (1979: 108). Cicindela (Cylindera) morio : FREITAG & BARNES (1989: 320). Cylindera (s. str.) morio : WIESNER (1992: 185), WIESNER (2020: 274).

Type localities. Cicindela morio : Brazil; Cicindela egena : Brésil: “les contrées riveraines du fleuve des Amazones [= the regions bordering the Amazon River]”.

Type material. Cicindela morio : HOLOTYPE: J ( MFNB), labelled: “3719” [printed] // “morio / Klg. / 307 / Brasil. Freir.” [large green collection label with thin black frame, handwritten] // “Hist. - Coll. ( Coleoptera ) / Nr 3719 / Cicindela morio Kl. / Brasil Freireiss / Zool Mus . Berlin” [green label with black frame, printed] // “ Holotype / Cicindela / morio / Klug, 1834 / labelled by MFNB 2024” [red label, printed] // “ Cylindera (s. str.) / morio ( Klug, 1834) / det. Jiří Moravec 2024” [printed].

Cicindela egena : LECTOTYPE (designated here): ♀ ( MNHN), lacking labels but standing as first along the large, ochre-tarnished collecting label with black frame: “egena / Chaud. / Amér. équin. / Amazones, / 52 Deyrolle” [handwritten] and labelled: “ Lectotype / Cicindela egena / Chaudoir, 1854 / design. Jiří Moravec 2024” [red label, printed] // “ Cylindera (s. str.) / morio ( Klug, 1834) / det. Jiří Moravec 2024” [printed]. Other material examined. HISTORICAL SPECIMENS: 1 J ( MNHN) [standing as C. egena along with the above-addressed lectotype of the synonymous taxon] with small, brownish, square green label. BRASIL: 2 JJ 1 ♀ ( MFNB): “ Ecuador ” [Equator] / Manaos”. 4 JJ 2 ♀♀ ( IRSNB), 1 J (CCJM, ex IRSNB): “ Brazil /Aurora” // “Coll. R.I. Sc. N. B. / Brazil ”. 1 J ( IRSNB): “802” / S. Paulo / Brasil ” // “Coll. J. Muller / Cicindela / morio / Kl. / R.M.H.N.B. 16.364”. 1 ♀ ( IRSNB): “ Brésil / Sao Paulo / Coll. Schramm” // “R. Mus . Hist. Nat. / Belg. I. G. 11. 230”. 1 J ( BMNH): “ Brazil ” // “obscurela Kl.”. 2 JJ 1 ♀ ( BMNH): “St. Paul” // “296 s.t.” // “F. Bates Coll. / 1911–248”. 1 ♀ ( BMNH): “Santarem / Lower Amazon / 28.I.96 / E. E.Austen / 96–80”. 1 ♀ ( BMNH): “ Brazil / Santarem” // “52/96” [on opposite side of the label] // “ C. morio / Klug” // “named by Dr. W. Horn / G.J.A.” // “F. Bates Coll. / 1911–248”. 1 J 1 ♀ ( BMNH): “Jatahy / Prov. Goyaz Brésil ”. 1 J 1 ♀ ( BMNH): same labels and: “F. Bates Coll. / 1911–248” // “morio / D. Klug” // “t. Horn” // “ Cicindela / morio Klug / det. R. Freitag 1984”. 1 ♀ ( SDEI): “Jatahy / Goyaz”. 1 J ( SDEI): “Manaos / IX. 27”. 2 JJ ( SDEI): “Staudinger / Sao Paulo ”. 1 ♀ ( HNHM): “Brasilia / S. Paulo” // “ C. morio Klug / det W.Horn”. 1 J ( SDEI): “ Brésil, Goyaz / Rio Verde”.RECENT SPECIMENS: 1 J ( MZSP): “Utiariti / Rio Papagaio, Mt. / XI.1996 / Lenko & Pereira” // “MZSP6228”. 1 ♀ ( MZSP):“Faz Cachoeirinha / Jataí, Goiás – Brasil / X.1962 / Exp. Dep. Zool.” // “MZSP62729”. 1 ♀ ( MZSP): “Faz.Acerio / Jataí Goiás / Brasil / X.1962 / Exp. Dep. Zool.” // “MZSP62731”. 1 J 1 ♀ ( JWCM): “ Brazil: Amazonas / 20 km n.e. Manaus / 18.IX.1992 / D. Pearson” // “clay-sand / savanna clearing”. BOLIVIA: 1 J ( COSJ): “ Bolivia – Santa Cruz department / Concepcion env. / 16°07´24´´S, 62°00´09´´W / 7-9.I. 2020, 494 m / O. Šafránek & M.Amaya lgt.”. 2 ♀♀ ( COSJ): “ Bolivia – Santa Cruz department / Concepcion env., 494 m / 16°07´24´´S, 62°00´09´´W / 10.I.2016 / O. Šafránek & J. L. Aramayo lgt.”. 1 J ( FCCR-MCZR): “ Bolivia – Santa Cruz / 25.8 km S / Rafael 305m / D. Brzoska 3-XII-1995 ”.

Examined specimens were labelled: “ Cylindera (s. str.) / morio ( Klug, 1834) / det. Jiří Moravec 2024” (“2025” respectively).

Differential diagnosis and brief redescription. Cylindera morio is principally distinguished from all other species of this group by its mandibles with four teeth (apart from basal molar) ( Figs 1 , 12 , 27–30, 32–33 , 35, 38 , 44, 46, 48 ), combined with distinct outer subapical lobe on terminal tooth of right male mandible in its lateral view ( Figs 2 , 13 , 31, 34 , 45, 47 ), very different shape of its aedeagus ( Figs 17–18 , 54–62 ) and generally much larger body size. Such distinct outer lobe on right male mandible (also visible when the mandibles are closed) is shared only with Cylindera amayai sp. nov., which, however, clearly differs in having tridentate mandibles ( Figs 3, 5 , 81, 83, 85, 87 ) and complex of other diagnostic characters, including its sparsely punctate-setose proepisterna ( Fig. 16 ) and shorter aedeagus apex ( Figs 54–62 ).

Body ( Figs 11 , 21 , 25–26 ) largest within the species-complex, (7.50–)7.90–10.2 (holotype 8.00) mm long, 2.70–3.40 (holotype 2.90) mm wide (lectotype of C. egena 9.20 mm long, 3.25 mm wide). Labrum ( Figs 14 , 27–29, 32–33 , 36–38, 40–43 , 49–51 ) yellow-testaceous, anterior margin semicircular in both sexes yet usually more distinctly in females, variably with 4–9 teeth which are blunt except for subacute median tooth (usually more protruding in females). Antennae as in others of the species complex, but antennomeres 5–6 mostly darker.

Thorax. Pronotum ( Figs 15–16 , 52–53 ) 1.70–2.15 mm long, 1.90–2.30 mm wide; surface of pronotal disc extremely finely asperate, finely irregularly wrinkled only at base and on pronotal posterior lobe, but lacking parallel striae; the fine pronotal sculpture is shared with Cylindera acompsa , which, however, has its mandibles with only three teeth (apart from basal molar) as also does Cylindera amayai sp. nov. which, moreover, has its pronotal surface mostly extremely finely micro-granulate. Proepisterna ( Fig. 16 ) distinctly foveolate-punctate-setose.

Elytra ( Figs 20 , 23–24 , 63–72 ) oblong, 5.20–6.50 mm long, outer margins subparallel in males, moderately dilated below middle in females, from their arcuate anteapical angles attenuated towards small sutural spine, thus forming elongate-acute apex (more conspicuously in males); surface almost black with darker velvety-black streaks and 5–7 foveae which may be greenish and variably conspicuous or barely recognizable; whitish elytral maculation either absent or with small or only indicated anteapical macula, exceptionally also lateromedian spot present ( Fig. 67 ).

Aedeagus ( Figs 17–18 , 54–62 ) elongate, 2.30–2.60 mm long, 0.50–0.55 mm wide, in its lateral view conically attenuated towards more or less elongate and variably (yet always noticeably) dorsally emarginate apex; the aedeagus apex in holotype is slightly damaged, unnoticeably crashed ventrally, yet somewhat changing its shape ( Fig. 17 ).

Distribution and biology. As mentioned under the redescription of the species-complex above, the distribution of Cylindera morio in literature includes also the other taxa of the species-complex. HORN (1915) mentioned “ Para bis Manaos, Matto Grosso, Goiás, Sao Paulo,?Nord-Argentina”, while later, apart from Lower and Upper Amazonia, HORN (1926) mentioned also the state of Minas Gerais. Although no specimen from Minas Gerais was recorded by FREITAG & BARNES (1989) and none was examined within the present revision, the occurrence in the state might be possible in the areas neighbouring with the state of Goiás, covered with savanna biotopes classified as Cerrado biome ( Fig. 214 ). NAVIAUX (2002) mentioned “ Sao Paulo, Mato Grosso (obviously including Mato Grosso do Sul), Goiás, Para and Amazon”. This species was recently recorded from the Bolivian department of Santa Cruz by GUERRA et al (1997), as well as by PEARSON et al. (1999) with a map of distribution.

The occurrence of Cylindera morio (and of any taxon of this species-complex) in Argentina is uncertain (see “Distribution” under “General redescription of the species-complex” above).

Consequently, only verified localities based on examination (either personally or from photos gained from colleagues mentioned in each of the data or illustrations) are listed here. As the labels of types and historical specimens mostly bear only the names of the Brazilian states without specified areas, the distribution on the map ( Fig. 215 ) is marked partly approximately. The “ Sao Paulo ” (in one case “S. Paul”) on the labels of several specimens in collections might at least partly mean São Paulo de Olivença situated on the western border of the Brazilian state of Amazonas, rather than the remote state of São Paulo lying in the southeastern coast with the Atlantic Rainforest (Mata Atlantica). However, the occurrence is possible in areas covered with savanna biotopes (Cerrado biome) as confirmed by one female in MZSP (not exactly examined here) recorded by FREITAG & BARNES (1989) from “Avanhand” which means Avanhandava in the western part of the state of São Paulo (see the map in Fig. 215 ); the area is now surrounded by plantations, thus missing its original biotopes (Gabriel Biffi (MZSP), pers. comm.).

In spite of the possible confusions of the taxa and their rare depository in collections (for instance only one in ZSM and HNHM, three in MCZR and three in NMPC), or absence (such as in NHMW), Cylindera morio is obviously rather widely distributed. It is obviously partly sympatric with others of this species-complex in Brazil. Records of this species from Manaos within the Amazon rainforest biome probably come from sandy-grassy areas near the riversides, yet probably not directly from sandy beaches. GUERRA et al. (1997) and PEARSON et al. (1999) mentioned that in Bolivia the species was found on light sand in open grassy areas in cerrado-savanna. This is also in accordance with the herein examined and illustrated specimens recently collected by the second author in the area of the city of Concepción in the Bolivian department of Santa Cruz, where Cylindera morio inhabits more open areas within the low forest vegetation ( Fig. 219 ). According to the ecoregion delineation by IBISCH et al. (2004), the locality near Concepcion meets two ecoregions: Chiquitano dry forest and Cerrado forest of the Chiquitano region. Observed adults of this species run and fly quickly along the paths during the day and fly quickly when disturbed, as also addressed by PEARSON et al. (1999).

Remarks. FREITAG & BARNES (1989) did not examine the type specimens of any taxon of the Cylindera morio species-complex, and failed to examine mandibles which are firmly closed in old specimens. Therefore, they overlooked the above-emphasized important diagnostic characters, despite the fact that one of the characters, the outer subapical lobe on the right male mandible in Cicindela (Cylindera) morio , is visible even when the mandibles are closed, and that the lobe is absent in other Brazilian taxa of the species-complex. Consequently, the redescription under Cicindela (Cylindera) morio and the map of distribution by FREITAG & BARNES (1989) includes also other Brazilian species of the complex (see also “Remarks” under Cicindela (Cylindera) acompsa below).

Figures

Figs 1–10. Cylindera (Cylindera) morio species-complex, diagnostic characters on male mandibles. 1–2 – C. (C.) morio (Klug, 1834): 1 – with four teeth; 2 – terminal tooth of right mandible in lateral view. 3–6 – C. (C.) amayai sp. nov.: 3 – with three teeth; 4 – terminal tooth of right mandible in lateral view; 5 – with small additional tooth in right mandible; 6 – terminal tooth of right mandible in lateral view. 7–10 – C. (C.) acompsa (Chaudoir, 1852) (and other two species of the complex): 7 – with tree teeth; 8 – terminal tooth of right mandible in lateral view; 9 – with three teeth and additional rudiment of tooth; 10 – terminal tooth of right mandible in lateral view (its outer margin in the three later species is smooth also in mandibles possessing additional small tooth). Scale bars = 1 mm.

Figs 11–20. Cylindera (Cylindera) morio (Klug, 1834), holotype, J (MFNB). 11 – habitus, 8 mm; 12 – head part; 13 – terminal tooth of right mandible in lateral view (its apical half); 14 – labrum; 15 – pronotum, dorsal view; 16 – pronotum, lateral view, showing proepisternum; 17–18 – aedeagus (apical portion) in its left lateral and dorsal aspects; 19 – labels; 20 – elytron. Scale bars = 1 mm.

Figs 21–24. Cylindera (Cylindera) morio (Klug, 1834). 21–23 – ♀, lectotype (MNHN) of the synonym Cicindela egena Chaudoir, 1854: 21 – habitus, 9.2 mm; 22 – labels; 23 – elytron. 24 – male from collection Chaudoir (MNHN), elytron. Scale bars = 1 mm.

Figs 25–26. Cylindera (Cylindera) morio (Klug, 1834), habitus. 25 – J, 8.8 mm, Brazil, Aurora (IRSNB); 26 – J, 8.4 mm, Bolivia, Concepción (COSJ). Scale bar = 1 mm.

Figs 27–34. Cylindera (Cylindera) morio (Klug, 1834). 27 – head, J, Brazil,Aurora (IRSNB); 28 – mandibles with labrum, ♀, Brazil, Santarem (BMNH); 29 – right mandible with labrum, ♀, Brazil, Jataí (MZSP 62731); 30 – mandibles, J, Brazil, “Goyaz” [= Goiás] (SDEI); 31 – J, Brazil, “Goyaz”, right mandible in lateral view; 32–33 – mandibles with labrum, Brazil, São Paulo: 32 – ♀ (BMNH); 33 – J (SDEI); 34 – J, Brazil, São Paulo, terminal tooth of right mandible in lateral view. Scale bars = 1 mm. (For Fig. 29 image credit Gabriel Biffi).

Figs 35–43. Cylindera (Cylindera) morio (Klug, 1834). 35–36 – ♀, lectotype (MNHN) of the synonym Cicindela egena Chaudoir, 1854: 35 – head; 36 – labrum; 37–39 – J of the synonym C. egena (MNHN): 37 – labrum; 38 – mandibles; 39 – J, terminal tooth of right mandible in lateral view; 40–43 – labrum: 40–42 – Brazil, Aurora: 40–41 – J (IRSNB); 42 – J (CCJM ex IRSNB); 43 – J, “Manaos” [= Manaus] (MFNB). Scale bars = 1 mm.

Figs 44–51. Cylindera (Cylindera) morio (Klug, 1834). 44 – mandibles, J, “Manaos” [= Manaus] (MFNB); 45 – J, terminal tooth of right mandible in lateral view; 46–48 – head, Bolivia, Concepción (COSJ): 46 – J; 47 – J, terminal tooth of right mandible in lateral view; 48 – ♀; 49–51 – labrum, Bolivia, Concepción (COSJ): 49 – J; 50–51 – ♀. Scale bars = 1 mm.

Figs 52–62. Cylindera (Cylindera) morio (Klug, 1834). 52–53 – pronotum: 52 – J, Brazil, Aurora (IRSNB); 53 – J, Bolivia, Concepción (COSJ); 54–62 – male aedeagus or its apex: 54 – Brazil, Aurora (IRSNB); 55–56 – Brazil, Aurora, internal sac in its left and right lateral aspect; 57–58 – Brazil, Aurora (IRSNB); 59 – Brazil, São Paulo (BMNH); 60 – Brazil, São Paulo (IRSNB); 61 – Bolivia, Concepción (COSJ); 62 – Brazil, Manaus (MFNB). Scale bars = 1 mm.

Figs 63–72. Cylindera (Cylindera) morio (Klug, 1834), elytron. 63–66 – Brazil,Aurora (IRSNB): 63–65 – J; 66 – ♀; 67 – J, São Paulo (SDEI); 68 – J, Manaus (MFNB); 69 – J, Goiás (SDEI); 70–72 – Bolivia, Concepción (COSJ): 70 – J; 71–72 – ♀. Scale bar = 1 mm.

Figs 81–97. Cylindera (Cylindera) amayai sp. nov., all from Bolivia, Palmarito. 81 – head, J, holotype (UASC, temporarily in COSJ); 82 – J, holotype, terminal tooth of right mandible in lateral view; 83–86 – mandibles, J, paratypes (COSJ): 83 and 85 – both mandibles; 84 and 86 – terminal tooth of right mandible in lateral view; 87 – head, ♀, allotype (COSJ); 88–91 – labrum: 88–90 – J, paratypes; 91 – ♀, allotype; 92–97 – male aedeagus: 92 – holotype; 93 – paratype; 94 – paratype, dorsal view; 95–96 – internal sac in left and right lateral views; 97 – paratype, apex. Scale bars = 1 mm.

Fig.214.Biomes in Brazil.(Approximately compiled according to a map by Toda Matéria https://www.todamateria.com.br/biomas-brasileiros/).

Fig. 215. Distribution of the Cylindera morio species-complex in Brazil and Bolivia, verified records (modified from the map by QGIS Development Team, 2016).

Fig. 219. Biotope of Cylindera morio Klug, 1834, near Concepción, Bolivia.