Ovipositor and mouthparts in a fossil insect support a novel ecological role for early orthopterans in 300 million years old forests
- College of Life Sciences and Academy for Multidisciplinary Studies, Capital Normal University, China
- Institute of Ecological Agriculture, College of Agronomy, Sichuan Agricultural University, China
- School of Life Sciences, Guangzhou University, 230 Waihuanxi Road, Guangzhou Higher Education Mega Center, China
- Institute of Evolutionary Biology and Animal Ecology, University of Bonn, Germany
- CR2P (Centre de Recherche en Paléontologie – Paris), MNHN – CNRS – Sorbonne Université; Muséum National d’Histoire Naturelle, France
Figures
Figure 1
Ctenoptilus frequens sp. nov., holotype (CNU-NX1-326).
(A) Habitus drawing and (B) habitus photograph (composite); (C–D) details of head and right foreleg (location as indicated in B), (C) color-coded interpretative drawing and (D) photograph …
Figure 2
External ovipositor in Ctenoptilus frequens sp. nov. in lateral view.
(A–C) Specimen CNU-NX1-749, (A) overview of the ovipositor with overlaid indications of the ovipositor parts (see also) overview of the ovipositor with overlaid indications of the ovipositor parts …
Figure 3
The evolution of major ovipositor configurations across Orthoptera.
(A) External ovipositor of external ovipositor of Ceuthophilus sp. (Orthoptera: Rhaphidophoridae; extant species) in laterial view (left side, flipped horizontally, left gonostylus IX [gs9] …
Figure 4
Head morphology (A–D) in Ctenoptilus frequens sp. nov. and (E) mandibular mandibular mechanical advantage in Ct. frequens sp. nov. and a selection of polyneopteran species.
(A–B) Specimen CNU-NX1-754, (A) color-coded interpretative drawing, and (B) photograph (composite) (as located on Appendix 1—figure 7I); (C–D) Specimen CNU-NX1-764, (C) color-coded interpretative …
Figure 5
Reconstruction of a female of Ctenoptilus frequens sp. nov. laying eggs.
Courtesy of Xiaoran Zuo.
Appendix 1—figure 1
Workflow for the extraction of the mandibular mechanical advantage based on 3D models.
Appendix 1—figure 2
Ctenoptilus frequens sp. nov., specimens composed of fore- and hind wings in connection with body remains.
(A–B) Specimen CNU-NX1-752; habitus, left forewing as positive imprint and right forewing and hind wings as negative imprints, (A) drawing and (B) photograph (composite). (C–D) Specimen CNU-NX1-738; …
Appendix 1—figure 3
Ctenoptilus frequens sp. nov., specimens composed of fore- and hind wings in connection with body remains.
(A–B) Specimen CNU-NX1-759; habitus, left hind wing as positive imprint and right wings as negative imprints, (A) drawing (for clarity, drawing of right hind wing venation duplicated and relocated, …
Appendix 1—figure 4
Ctenoptilus frequens sp. nov., specimens composed of fore- and hind wings in connection with body remains.
(A–D) Specimen CNU-NX1-747; (A–B) habitus, all wings as negative imprints, (A) drawing (for clarity, drawing of right hind wing venation duplicated and relocated, original location in light grey on …
Appendix 1—figure 5
Ctenoptilus frequens sp. nov., specimens composed of forewings, isolated or by pair, and forewing and ovipositor.
(A–B) Specimen CNU-NX1-748; right forewing, negative imprint, (A) drawing and (B) photograph (composite, flipped horizontally, light-mirrored). (C–D) Specimen CNU-NX1-732; right forewing, positive …
Appendix 1—figure 6
Ctenoptilus frequens sp. nov., specimens composed of well-exposed hind wings in connection with body remains or isolated.
(A–B) Specimen CNU-NX1-198; (A) drawing of right hind wing (location as indicated in B) and (B) photograph of habitus (composite, flipped horizontally), left forewing as positive imprints and left …
Appendix 1—figure 7
Ctenoptilus frequens sp. nov., specimens composed of body remains including well-preserved head, legs, and/or ovipositor.
(A–E) Specimen CNU-NX1-749; (A) photograph of habitus (composite), left forewing as positive imprint; (B–C) details of ovipositor (location as indicated in A; to be compared with main document Figure…
Appendix 1—figure 8
Ctenoptilus frequens sp. nov., specimen CNU-NX1-742.
(A) Photograph of habitus (composite), right forewing as positive imprint, flipped horizontally, and (B–C) details of ovipositor (location indicated in A; to be compared with main document). …
Appendix 1—figure 9
Outlines of the mandibular gnathal edges for all studied taxa.
Appendix 1—figure 10
Progression of mechanical advantage curves for the studied taxa.
x-axis = % tooth row; y-axis = MA (mechanical advantage).
Appendix 1—figure 11
Results of the principal component (PC) analysis of the mandibular mechanical advantage for the first two PCs together with results for the first two PCs after phylogenetic signal correction.
Large dots, distribution of species in PC space uncorrected for phylogenetic signal; small dots, distribution of species in PC space corrected for phylogenetic signal. Although phylogenetic signal …
Appendix 1—figure 12
Correspondence between terminologies applied to polyneopteran insect ovipositors.
Tables
Appendix 1—table 1
Food preference of polyneopteran species included in the mechanical advantage (MA) principal component analyses.
| Order | Species | Food preference |
|---|---|---|
| Dermaptera | Diplatys flavicollis | Omnivore |
| Dermaptera | Forficula auricularia | Omnivore |
| Dermaptera | Hemimerus sp. | Carnivore |
| Embioptera | Aposthonia japonica | Herbivore |
| Embioptera | Embia ramburi | Herbivore |
| Embioptera | Metoligotoma sp. | Herbivore |
| Grylloblattodea | Grylloblatta bifratrilecta | Omnivore |
| Orthoptera | Acheta domesticus | Omnivore |
| Orthoptera | Comicus calcaris | Omnivore |
| Orthoptera | Conocephalus sp. | Omnivore |
| Orthoptera | Myrmecophilus sp. | Omnivore |
| Orthoptera | Gryllus bimaculatus | Omnivore |
| Orthoptera | Cyphoderris sp. | Omnivore |
| Orthoptera | Hemideina crassidens | Omnivore |
| Orthoptera | Meconema meridionale | Carnivore |
| Orthoptera | Papuaistus sp. | Omnivore |
| Orthoptera | Prosopogryllacris sp. | Omnivore |
| Orthoptera | Stenobothrus lineatus | Herbivore |
| Orthoptera | Stenopelmatus sp. | Omnivore |
| Orthoptera | Pholidoptera griseoaptera | Omnivore |
| Orthoptera | Tettigonia viridissima | Omnivore |
| Orthoptera | Tridactylus sp. | Herbivore |
| Orthoptera | Troglophilus neglectus | Omnivore |
| Orthoptera | Xya variegata | Fetritivore |
| Phasmatodea | Agathemera sp. | Herbivore |
| Phasmatodea | Peruphasma schultei | Herbivore |
| Plecoptera | Eusthenia lacustris | Carnivore |
| Plecoptera | Perla marginata | Carnivore |
| Zoraptera | Zorotypus caudelli | Herbivore |
| Orthoptera | Ctenoptilus frequens CFMR | Tested |
| Orthoptera | Ametrosomus sp. | Omnivore |
| Orthoptera | Ametrus tibialis | Omnivore |
| Orthoptera | Apotrechus illawarra | Omnivore |
| Orthoptera | Bothriogryllacris brevicauda | Omnivore |
| Orthoptera | Chauliogryllacris grahami | Omnivore |
| Orthoptera | Cnemotettix bifascicatus | Omnivore |
| Orthoptera | Cooraboorama canberrae | Omnivore |
| Orthoptera | Kinemania ambulans | Omnivore |
| Orthoptera | Mooracra sp. | Omnivore |
| Orthoptera | Nullanullia maitlia | Omnivore |
| Orthoptera | Nunkeria brochis | Omnivore |
| Orthoptera | Paragryllacris combusta | Omnivore |
| Orthoptera | Pararemus sp. | Omnivore |
| Orthoptera | Wirritina brevipes | Omnivore |
Appendix 1—table 2
Importance and factor loadings of the principal component analyses of the polynomial regressions of the mechanical advantages (MAs).
| Principal component analysis | ||||||
|---|---|---|---|---|---|---|
| PC1 | PC2 | PC3 | PC4 | PC5 | PC6 | |
| Standard deviation | 0.380 | 0.158 | 0.083 | 0.061 | 0.039 | 0.025 |
| Proportion of variance | 0.794 | 0.136 | 0.038 | 0.021 | 0.008 | 0.003 |
| Cumulative proportion | 0.794 | 0.930 | 0.968 | 0.988 | 0.997 | 1.000 |
| Factor loadings: | ||||||
| PC1 | PC2 | PC3 | PC4 | PC5 | PC6 | |
| Intercept | −0.013 | 0.288 | 0.150 | 0.907 | 0.258 | 0.074 |
| Regression coefficient 1 | −0.948 | −0.250 | 0.173 | 0.046 | −0.048 | 0.065 |
| Regression coefficient 2 | 0.300 | −0.754 | 0.497 | 0.090 | 0.186 | 0.229 |
| Regression coefficient 3 | −−.037 | 0.509 | 0.528 | −0.372 | 0.290 | 0.489 |
| Regression coefficient 4 | −0.057 | −0.164 | −0.650 | −0.015 | 0.436 | 0.597 |
| Regression coefficient 5 | 0.079 | 0.028 | −0.010 | 0.171 | −0.789 | 0.585 |
| Phylogenetic principal component analysis | ||||||
| PC1 | PC2 | PC3 | PC4 | PC5 | PC6 | |
| Standard deviation | 0.745 | 0.368 | 0.172 | 0.131 | 0.099 | 0.055 |
| Proportion of variance | 0.740 | 0.180 | 0.040 | 0.023 | 0.013 | 0.004 |
| Cumulative proportion | 0.740 | 0.921 | 0.960 | 0.983 | 0.996 | 1.000 |
| Factor loadings: | ||||||
| PC1 | PC2 | PC3 | PC4 | PC5 | PC6 | |
| Intercept | 1.000 | −0.021 | −0.003 | 0.003 | 0.000 | 0.000 |
| Regression coefficient 1 | −0.088 | −0.995 | −0.048 | 0.009 | 0.005 | 0.004 |
| Regression coefficient 2 | 0.054 | −0.228 | 0.940 | −0.247 | −0.008 | −0.024 |
| Regression coefficient 3 | 0.076 | 0.026 | −0.412 | −0.905 | 0.067 | −0.006 |
| Regression coefficient 4 | −0.023 | 0.067 | 0.096 | 0.101 | 0.985 | 0.078 |
| Regression coefficient 5 | 0.077 | 0.085 | 0.180 | −0.112 | −0.241 | 0.940 |
