Kalligramma haeckeli Walther, 1904

Makarkin, Vladimir N. & Ansorge, Jörg, 2025, Redescription of Kalligramma haeckeli Walther, 1904 (Neuroptera: Kalligrammatidae), a Jurassic paleontological icon, allows the redefinition of Kalligrammatinae, Zootaxa 5659 (1), pp. 69-86 : 72-76

Kalligramma haeckeli Walther, 1904

Figs 3–9

Kalligramma Haeckeli Walther, 1904: 184 ; Pl. 8 [original description; photograph of the holotype]; Handlirsch 1906 –1908 [1906]: 611; Pl. 48, Fig. 12 [redescription; reconstruction of the fore- and hind wing venation]; Wannach 1910: 159, Pl. 5, Fig. 14 [reconstruction of the fore- and hind wing venation by Handlirsch (1906 –1908)]; Handlirsch 1911: 183; Pl. 10, Fig. 17 [reconstruction of the holotype]; Zittel 1915: Fig. 1448 [photograph of the holotype]; Bölsche 1916: 69, Fig. 10 [photograph of the holotype]; Tornquist 1916: 225, Fig. 68 [photograph of the holotype]; Abel 1917: 52, 53, Figs 25, 26 [photograph of the holotype and its reconstruction by Handlirsch (1911)]; Handlirsch 1919: 61, 62; Abel 1920: 128, Fig. 183 [reconstruction of the holotype by Handlirsch (1911)]; Handlirsch 1920 –1921 [1920]: 196, Fig. 163 [reconstruction of the holotype by Handlirsch (1911)]; Zittel 1921: Fig. 1447 [photograph of the holotype]; Abel 1922: 514, 515, 517, Fig. 425 [reconstruction of the holotype by Handlirsch (1911)]; Schaffer 1924: 337, Fig. 410 [reconstruction of the holotype by Handlirsch (1911)]; Zittel 1924: Fig. 1456 [photograph of the holotype]; Abel 1926: Fig. 17 [reconstruction of the holotype by Handlirsch (1911)]; Bölsche 1931: 154, Fig. 72 [simplified reconstruction of the holotype by Handlirsch (1911)]; Wagner 1931: 473, Bild 390 [reconstruction of the holotype by Handlirsch (1911)]; Jeannel 1946: 60, Pl. 11, Fig. 64 [reconstruction of the holotype by Handlirsch (1911)].

Kalligramma haeckeli : Paoli 1913: 38, Fig. 24 [reconstruction of the holotype by Handlirsch (1911)]; Berlese 1925: 172, Fig. 177 [reconstruction of the holotype by Handlirsch (1911)]; Cockerell 1928: 128; Hennig 1932: 422, Fig. 195 [reconstruction of the holotype by Handlirsch (1911)]; Handlirsch & Beier 1936: 1454, Fig. 1522 [reconstruction of the holotype by Handlirsch (1911)]; Martynova 1947: 2055–2057; Grassé 1949: 70, Fig. 73 [reconstruction of the holotype by Handlirsch (1911)]; Martynova 1949: 167; Rohdendorf 1949: 84, Fig. 75 [reconstruction of the holotype by Handlirsch (1911)]; Laurentiaux 1953: 480; Brues et al. 1954: 816, Fig. 1186 [reconstruction of the forewing venation by Handlirsch (1906 – 1908)]; Lotze 1955: Fig. 63 [reconstruction of the holotype by Handlirsch (1911)]; Shvanvich 1959: Fig. 175 [reconstruction of the holotype by Handlirsch (1911)]; Martynova 1962: Fig. 865 [reconstruction of the fore- and hind wing venation by Handlirsch (1906 –1908)]; Kéler 1963: Fig. 158 [reconstruction of the holotype by Handlirsch (1911)]; Kuhn 1963: Fig. 42 [reconstruction of the holotype by Handlirsch (1911)]; Müller 1963: 218, Fig. 298 [reconstruction of the holotype by Handlirsch (1911)]; Panfilov 1968: 166–170; Hennig 1981: 386; Jarzembowski 1984: Fig. 33 [simplified reconstruction of the holotype by Handlirsch (1911)]; Wellnhofer 1988: 88 [photograph of the holotype]; New 1989: 3, Figs 1 [reconstruction of the holotype by Handlirsch (1911)], 2 [reconstruction of the fore- and hindwing venation by Handlirsch (1906 –1908)]; Barthel et al. 1990: Fig. 7.46 [photograph of the holotype]; Carpenter 1992: 351, Fig. 195.4 [reconstruction of the fore- and hind wing venation by Handlirsch (1906 –1908)]; Seibold 1992: Front cover [reconstruction of the holotype by Handlirsch (1911)]; Frickhinger 1994: 154, Fig. 308 [photograph of the holotype]; Rumbucher 1995: Fig. 5 [reconstruction of the holotype by Handlirsch (1911)]; Ren & Guo 1996: 465; Mayr 2000: 146–151, Figs 1 , 2 [photographs of the holotype], Fig. 7 [reconstruction of the holotype by Handlirsch (1911)]; Andersen 2001: 434; Jarzembowski 2001: 56, 57; Zhang & Zhang 2003: 301, 303, 305; Scheven 2004: Fig. on p. 71 [photograph of the holotype]; Liu et al. 2014: 66, 68; Yang et al. 2014a: 918, 919; Yang et al. 2014b: 2 (Table), 11; Liu et al. 2015a: 339, 441; Khramov et al. 2020: 896.

Calligramma [sic] haeckeli : Franz 1924: 544, Fig. 537 [reconstruction of the holotype by Handlirsch (1911)].

Kalligramma Häckeli : Endriss 1927: 383, Fig. 563 [photograph of the holotype]

Kalligramma : Dacqué 1936: 129, Fig. 87 [photograph of the holotype].

Holotype. BSPG 1902 I 6 (part, counterpart), deposited in Bayerische Staatssammlung für Paläontologie und Geologie, München [=Munich], Germany [Paläontologisches Museum München].An incomplete specimen missing the left forewing and all legs; body very poorly preserved, partly hidden by the right hind wing. Solnhofener Bruch [= Solnhofen quarry] southwest of Solnhofen, Bavaria, Germany; Late Jurassic (early Thitonian), Weissjura- Gruppe, Altmühltal Formation ( Niebuhr 2015), Solnhofen Subformation, Malm Zeta 2 a/b ( Quenstedt 1856 –1857), Hybonotum-Zone (ca. 150 Ma, Hesselbo et al. 2020).

Verbatim label data: (1) [original label] [typed] Palaeontologisches Museum / no [handwritten] 1902 I 6 / Kalligramma Haeckeli / Walther / Lithograph. Schiefer / [typed] Von / [handwritten] Solnhofen/ Orig. Walther Haeckel Festschrift / [typed] München; (2) [label of 1913 to 1934] [typed] No. 1902. I. 6 [red square glued to the label] 1307 / Kalligramma Haeckeli. Walther / Lithographischer Schiefer./ Solnhofen/ [handwritten] Holotyp.[us] u.[nd] [typed] Orig.[inal] z.[u] Walther Haeckel Festschrift; (3) [lable of Franz Xaver Mayr, 1934] [typed] Staatl.[iche] Samml.[ung] f[ür]. Paläontol.[ogie] u.[nd] Hist.[orische] Geologie. / No. [handwritten by Franz Xaver Mayr] Nach Grimm:/ Solnhofener Bruch / b.[ei] Solnhofen/ F.M. [= Franz Xaver Mayr] 1934 / [typed] München; (4) [probably latest label] [typed] Bayer.[ische]. Staatssammlung f[ür]. Paläontologie u[nd]. histor[ische]. Geologie München/ ( BSP)/ No. [handwritten] 1902 I 6 / Kalligramma Haeckeli WALTHER / Untertithon. Solnhofener Plattenkalke / Solnhofen/ Holotypus + Original zu Walther 1904 / s. 184, Taf. 8.

Diagnosis. Large species, with hind wing ca. 100 mm long [smaller, with hind wing less than 100 mm (except K. jurarchegonium )]. Forewing: Cubital space (between CuA, CuP) very narrow [moderately broad, at least basally in other species]. Hind wing: CuP deeply forked [shallowly forked in K. jurarchegonium , K. sharovi , K. liaoningense ].

Description. Head ca. 5 mm wide (in dorsal view), with two long (ca. 10 mm), slender elements of mouthparts (probably maxillary or labial palps). Mesonotum ca. 10 mm wide, with large mesoscutum ( Fig. 4 ). Abdomen ca. 40 mm long, details not discernible.

Right forewing broadly oval, 113 mm long as preserved (estimated complete length ca. 115 mm), ca. 51 mm wide. Costal space moderately broad, slightly narrowed towards fusion of Sc and RA, then dilated distally. All preserved subcostal veinlets forked: in basal part deeply forked, but not forked near margin; in middle and distal parts shallowly forked with two to four short branches (often these branches forked again near margin). Costal crossveins numerous, not arranged to gradate series. Subcostal space relatively broad throughout entire length, with dense crossveins. Sc fused distally RA; Sc+RA with few branches (poorly preserved). RA space slightly narrower than subcostal space in proximal half, nearly equal in distal half. RP with 12 branches (eleven branches proximad fusion of Sc and RA). RP1 dichotomously forked (most deeply forked comparing with other branches of RP); RP2–RP9 distally dichotomously forked; distal parts of RP10–RP12 not preserved. Basal part of M not preserved. MA strongly concave, thinner in proximal part then RP1 and MP1, distally dichotomously forked. Posterior trace of MP with four preserved anteriorly-directed branches. MP1 pectinately branched, with four dichotomously forked branches; MP2–MP4 dichotomously forked distally. Cu dividing into CuA, CuP very close to wing base (this part not preserved); both nearly parallel and close each other for most length. CuA very poorly, fragmentarily preserved, strongly convex; close to CuP through entire length. CuP concave, probably dichotomously forked distally (this part incompletely preserved). A1 very long, with four to five distal pectinate branches (partially not preserved), which forked one to three times. A2 long, strongly pectinately forked, with nine branches: five proximal branches dichotomously forked; sixth branch twice forked; seventh and eighth once forked; ninth simple. A3 fragmentarily preserved, short. Trichosors not detected, probably absent. Eyespot very poorly preserved; only rounded concave structure (ca. 6 mm in diameter) prominent (central pigmented disc? swelling?).

The left forewing was not detected during early preparation attempts ( Walther 1904, p. 184) and appears not to be preserved.

Hind wing broadly triangular, 93 mm long as preserved (left wing) (estimated complete length ca. 100 mm), ca. 69 mm wide. Costal space moderately broad, strongly dilated distally (its basal-most part not preserved). Basal subcostal veinlets not forked, other forked: shallowly forked once to twice in proximal half; dichotomously forked in distal part. Costal crossveins numerous, not arranged to gradate series. Subcostal space moderately broad, with dense crossveins. Sc fused distally RA; Sc+RA dichotomously forked. RA space slightly narrower than subcostal space, with dense crossveins. RP originates close to wing base (right wing), with 11 branches (of these, ten branches proximad fusion of Sc and RA) in left wing. RP1 originates at RP, deeply dichotomously forked. RP 2 in left wing incomplete, forked proximad RP1 fork; its anterior branch fused with RP3, posterior branch not fused with other branches. RP3 to RP9 similarly configured, deeply (to varying degree) dichotomously forked. RP10, RP11 twice forked distally. M forked basally. MA strongly concave, dichotomously forked distally. Posterior trace of MP nearly straight, with five anteriorly-directed branches (right wing). MP1 pectinately branched, with four (left wing) to five (right wing) dichotomously forked branches; MP2–MP5 dichotomously forked. Cross-venation in radial, radiomedial, medial spaces very dense, becoming rarer distally. Fork of Cu not preserved, probably near wing base. CuA rather close to MP, distally pectinately branched with four/five dichotomously forked branches. CuP forked very deeply; anterior branch pectinate, with three dichotomously forked branches; posterior branch dichotomously forked distally. A1 long, close to posterior branch of CuP, dichotomously forked distally. A2 long, pectinately branched, with eight dichotomously forked branches. A3 short, probably dichotomously forked (incompletely preserved). Trichosors not detected, probably absent. Eyespot poorly preserved; presumed central pigmented disc ca. 13 mm in diameter; two elongate-oval embossed swellings ca. 6 mm long.

Figures

FIGURE 1. Kalligramma haeckeli Walther, 1904. A, reconstruction of the holotype wings by Handlirsch (1906–1908, Pl. 48, Fig. 12); B, reconstruction of the holotype specimen by Handlirsch (1911, pl. 10, Fig. 17).

FIGURE 2. Simplified early Tithonian palaeogeography of Central Europe. Red star indicates the Solnhofen area; M, Munich; B, Berlin; N, Nürnberg; NSB, North Sea Basin; PB, Polish Basin; SGB, South German Basin (based mainly on Meyer & Schmidt Kaler 1989 and Ziegler 1990).

FIGURE 3. Kalligramma haeckeli Walther, 1904, holotype BSPG 1902 I 6. A, part; B, counterpart. Scale bars = 5 cm.

FIGURE 4. Preserved part of the body of Kalligramma haeckeli Walther, 1904, holotype BSPG 1902 I 6. h, head; mp, mouthparts; mst, mesothorax. Scale bars = 1 cm.

FIGURE 5. Right forewing of Kalligramma haeckeli Walther, 1904, holotype BSPG 1902 I 6.A, part; B, counterpart (converted to standard right view). Scale bar = 1 cm (both to same scale).

FIGURE 6. Drawing of the right forewing of Kalligramma haeckeli Walther, 1904, holotype BSPG 1902 I 6. Scale bar = 1 cm.

FIGURE 7. Right hind wingof Kalligramma haeckeli Walther, 1904, holotype BSPG 1902I 6.A, part; B, counterpart (converted to standard right view). Scale bar = 1 cm (both to same scale).

FIGURE 8. Left hind wing of Kalligramma haeckeli Walther, 1904, holotype BSPG 1902 I 6. A, part (converted to standard right view); B, counterpart. Scale bar = 1 cm (both to same scale).

FIGURE 9. Drawing of the hind wings of Kalligramma haeckeli Walther, 1904, holotype BSPG 1902 I 6.A, left wing (converted to standard right view); B, right wing. Scale bar = 1 cm (both to same scale).

FIGURE 10. Fragment of the right hind wing of Sophogramma lii Yang et al., 2009 (holotype CNU-NEU-LB2008001) showing the configuration of cubital veins. Scale bar = 5 mm.