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. 2014 Jun 9:14:126.
doi: 10.1186/1471-2148-14-126.

Mesozoic lacewings from China provide phylogenetic insight into evolution of the Kalligrammatidae (Neuroptera)

Qiang Yang, Yongjie Wang  1 , Conrad C Labandeira, Chungkun Shih, Dong Ren
Affiliations

Mesozoic lacewings from China provide phylogenetic insight into evolution of the Kalligrammatidae (Neuroptera)

Qiang Yang et al. BMC Evol Biol. .

Abstract

Background: The Kalligrammatidae are distinctive, large, conspicuous, lacewings found in Eurasia from the Middle Jurassic to mid Early Cretaceous. Because of incomplete and often inadequate fossil preservation, an absence of detailed morphology, unclear relationships, and unknown evolutionary trends, the Kalligrammatidae are poorly understood.

Results: We describe three new subfamilies, four new genera, twelve new species and four unassigned species from the late Middle Jurassic Jiulongshan and mid Early Cretaceous Yixian Formations of China. These kalligrammatid taxa exhibit diverse morphological characters, such as mandibulate mouthparts in one major clade and siphonate mouthparts in the remaining four major clades, the presence or absence of a variety of distinctive wing markings such as stripes, wing spots and eyespots, as well as multiple major wing shapes. Based on phylogenetic analyses, the Kalligrammatidae are divided into five principal clades: Kalligrammatinae Handlirsch, 1906, Kallihemerobiinae Ren & Engel, 2008, Meioneurinae subfam. nov., Oregrammatinae subfam. nov. and Sophogrammatinae subfam. nov., each of which is accorded subfamily-level status. Our results show significant morphological and evolutionary differentiation of the Kalligrammatidae family during a 40 million-year-interval of the mid Mesozoic.

Conclusion: A new phylogeny and classification of five subfamilies and their constituent genera is proposed for the Kalligrammatidae. These diverse, yet highly specialized taxa from northeastern China suggest that eastern Eurasia likely was an important diversification center for the Kalligrammatidae. Kalligrammatids possess an extraordinary morphological breadth and panoply of adaptations during the mid-Mesozoic that highlight our conclusion that their evolutionary biology is much more complex than heretofore realized.

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Figures

Figure 1
Figure 1
Images and camera lucida drawings of Affinigramma myrioneura sp. nov. A, B, C, Holotype CNU-NEU-NN2009-006P/C; D, E, F, Paratype CNU-NEU-NN2009-007P/C. Scale bars, 10 mm.
Figure 2
Figure 2
Images and camera lucida drawings of Kallihemerobius aciedentatus sp. nov. CNU-NEU-NN2010-008P/C; A, B, Holotype; C, Outline of holotype; D, Outline of forewing. Scale bars, 10 mm.
Figure 3
Figure 3
Images and camera lucida drawing of Kallihemerobius almacellus sp. nov. CNU-NEU-NN2009-050P/C. A, B, Holotype; C, Outline of holotype. Scale bars, 10 mm.
Figure 4
Figure 4
Images and camera lucida drawing of Kallihemerobius feroculus sp. nov. CNU-NEU-NN2010-013P/C. A, B, Holotype; C, Outline of holotype. Scale bars, 10 mm.
Figure 5
Figure 5
Images and camera lucida drawing of Stelligramma allochroma sp. nov. CNU-NEU-NN2010-012P/C. A, B, Holotype; C, Outline of holotype. Scale bars, 10 mm.
Figure 6
Figure 6
Image and camera lucida drawings of Kallihemerobiinae gen. et sp. indet. CNU-NEU-NN2009-033. A, Habitus; B, Outline of habitus. Scale bars, 10 mm.
Figure 7
Figure 7
Images and camera lucida drawing of Kalligramma brachyrhyncha sp. nov. CNU-NEU-NN2009-030P/C. A, B, Holotype; C, Outline of holotype. Scale bars, 10 mm.
Figure 8
Figure 8
Images and camera lucida drawings of Kalligramma circularia sp. nov. A, Holotype CNU-NEU-NN2010-003; B, Paratype CNU-NEU-NN2010-011; C, D, Paratype CNU-NEU-NN2010-015P/C; E, Outline of holotype; F, Forewing of holotype; G, Hind wing of holotype; H, Outline of paratype CNU-NEU-NN2010-011. Scale bars, 10 mm.
Figure 9
Figure 9
Images and camera lucida drawing of Kalligramma sp. CNU-NEU-NN2010-010P. A, B, Habitus; C, Outline of habitus. Scale bars, 10 mm.
Figure 10
Figure 10
Images and camera lucida drawing of Abrigramma calophleba sp. nov. CNU-NEU- HP2009-001P. A, B, Holotype; C, Outline of holotype. Scale bars, 10 mm.
Figure 11
Figure 11
Images and camera lucida drawing of Ithigramma multinervia sp. nov. CNU-NEU-NN2009-034P. A, B, Holotype; C, Outline of holotype. Scale bars, 10 mm.
Figure 12
Figure 12
Images and camera lucida drawing of Ithigramma sp. CNU-NEU-NN2010-016P. A, B, Habitus; C, Outline of habitus. Scale bars, 10 mm.
Figure 13
Figure 13
Images and camera lucida drawing of Oregramma aureolosa sp. nov. CNU-NEU- NN2009-032P. A, B, Holotype; C, Outline of holotype. Scale bars, 10 mm.
Figure 14
Figure 14
Images and camera lucida drawings of Oregramma illecebrosa sp. nov. CNU-NEU-LB2009-031C. A, B, Holotype; C, Outline of forewing with body; D, Outline of hind wing. Scale bars, 10 mm.
Figure 15
Figure 15
Images and camera lucida drawing of Oregramma sp. CNU-NEU-NN2010-014P. A, B, Habitus; C, Outline of habitus. Scale bars, 10 mm.
Figure 16
Figure 16
Images and camera lucida drawings of Sophogramma pingquanica sp. nov. CNU-NEU-HP2010-009P. A, B, Holotype; C, Outline of Holotype; D, Outline of forewing. Scale bars, 10 mm.
Figure 17
Figure 17
A composite system of the phylogenetic relationships within the Kalligrammatidae. 1: a, Parallel MP vein branches, b, Eyespot absence, c, Presence of a humeral recurrent vein (Vr). 2: a, A large triangular MP region, b, Presence wing eyespots. 3: Simple costal crossveins, with few interlinked veinlets. 4: a, Complex costal cross-venation, b, Deep MA forks. 5: Distal branching of MP. 6: a, Distal constriction of the costal region; b, Sinuate MP; c, Expanded region between CuA and CuP. 7: a, Broadly triangular forewing; b, 1A bifurcation at wing midsection or distally. 8: Proximal bifurcation of 1A vein. 9: a, Presence of a broad costal region; b, Few Rs branches. 10: Presence of an ORB.
Figure 18
Figure 18
Histograms of kalligrammatid species and generic diversity during their 40 million-year existence during the mid Mesozoic. A, Histogram showing the genus and species richness of the Kalligrammatidae. B, Histogram showing the genus and species richness at the subfamilial level within the Kalligrammatidae.
Figure 19
Figure 19
Structural diversity among Kalligrammatidae. A, Sophogramma lii Yang, Zhao and Ren, 2009. B, Oregramma aureolosa sp. nov. C, Stelligramma allochroma sp. nov. D, Affinigramma myrioneura sp. nov. E, Oregramma illecebrosa sp. nov. F, Kalligramma circularia sp. nov. G, Affinigramma myrioneura sp. nov. H, Apochrysogramma rotundum Yang, Makarkin and Ren, 2011. I, Kalligramma brachyrhyncha sp. nov.
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