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Fig. 8 in A general theory of genital homologies for the Hexapoda (Pancrustacea) derived from skeletomuscular correspondences, with emphasis on the Endopterygota

Fig. 8. Genital skeletomusculature of Hypera cf. postica (Coleoptera: Curculionidae). AeD, genitalia in situ within terminal abdominal segments: A, dorsal; B, lateral, left side, sternal-coxital muscle partially removed;C, ventral;D, lateral, right side. E, J, coxites I encircling coxites II and stylar composite,lateral,with sternum removed in E which shows the left side. FeJ, coxopenis (coxital-penial composite, CxtII þ Pen): F, endophallus partially exserted, ventrolateral oblique; G, dorsal; H, ventral; I, sternum in place, right side; J, ventrolateral oblique. Abbreviations: CxtI ¼ "phallobase" (first gonocoxites); CxtII þ Pen ¼ coxopenial composite; Endscl ¼ endophallic sclerite; StVIII ¼ sternum VIII; StIX ¼ sternum IX; Stap ¼ sternal apophysis; TgIX ¼ tergum IX. Muscle abbreviations indicated in Table 1.

Fig. 13 in A general theory of genital homologies for the Hexapoda (Pancrustacea) derived from skeletomuscular correspondences, with emphasis on the Endopterygota

Fig. 13. Genital skeletomusculature of Panorpa nuptialis (Mecoptera: Panorpidae). A, genitalia in situ, lateral. B, composite sclerite of tergum and sternum IX. CeE, genital appendages: C, dorsal; D, ventral; E, distal. FeH, right half of genital appendages: F, dorsolateral oblique; G, lateral, paired appendages of sperm pumping complex intact; H, lateral, left half of sperm pumping complex removed. I, J, mittelplatte and dorsal pseudoparameres (¼ kammersklerit plus second fragment of aedeagal apodem). K, pistilltrö ager, ventrolateral oblique.L, base of dorsal pseudoparameres, posterior view.M, base of dorsal pseudoparameres, internal (anterior) view. Abbreviations: Apd ¼ apodeme of pistilltrö ager; Ce ¼ cercus;d. ps. par. ¼ dorsal pseudoparamere; Cxa ¼ gonocoxa or first gonocoxites; Cxt ¼ gonocoxite; CxtAapd ¼ anterior apodeme of gonocoxite; CxtDapd ¼ dorsal apodeme of gonocoxite; mittelp. ¼ mittelplatte; pistillt. ¼ pistilltrö ager; Pen ¼ penis or penial sclerite; Prct ¼ proctiger; StIX ¼ sternum IX; Sty ¼ gonostylus; StyDcond ¼ dorsal stylar condyle; StyVcond ¼ ventral stylar condyle; TgIX ¼ tergum IX. Muscle abbreviations indicated in Table 1. Note: Sclerite fragment numbers indicated with Roman numerals; addition signs indicate sclerite fusion.

Fig. 10 in A general theory of genital homologies for the Hexapoda (Pancrustacea) derived from skeletomuscular correspondences, with emphasis on the Endopterygota

Fig. 10. Genital skeletomusculature of Mengenilla sp. (Strepsiptera: Mengenillidae), modified from Hünefeld, Pohl, et al. (2011b). A, abdominal segment IX and proctiger in cross-section. Abbreviations: Cxa ¼ gonocoxa; Pen ¼ penis; Prct ¼ proctiger; StIX ¼ sternum IX; Stap ¼ sternal apodeme; TgIX ¼ tergum IX. Muscle abbreviations indicated in Table 1.

Fig. 9 in A general theory of genital homologies for the Hexapoda (Pancrustacea) derived from skeletomuscular correspondences, with emphasis on the Endopterygota

Fig. 9. Genital skeletomusculature of Ilybius biguttulus (Coleoptera:Dytiscidae). AeD, genitalia in situ: A, dorsal; B, dorsolateral oblique; C, ventral; D, dorsal, tergal-coxital muscles removed.EeG, genitalia and anterior fragment of sternum IX (sternal apodeme): E, ventral; F, ventral,sternal muscle removed;G, dorsal.H, coxital-stylar composite sclerites spread away from coxopenis (composite of second coxites and penis, CxtII þ Pen), dorsolateral oblique. I, coxopenis, left side, lateral. Abbreviations: CxtI þ Sty ¼ coxostylar composite sclerite; CstII þ Pen ¼ coxopenial composite; Endscl ¼ endophallic sclerite; StIX ¼ sternum IX; Stap ¼ sternal apodeme; Styapd ¼ stylar apodeme; TgIX ¼ tergum IX. Muscle abbreviations indicated in Table 1.

Fig. 7 in A general theory of genital homologies for the Hexapoda (Pancrustacea) derived from skeletomuscular correspondences, with emphasis on the Endopterygota

Fig. 7. Genital skeletomusculature of Cantharis (Cantharis) (Coleoptera: Cantharidae). A, B, genitalia in situ: A, lateral; B, ventral. CeN, genitalia various aspects and states of dissection: C, whole, dorsal; D, E, stylus and muscle partially removed, dorsal; F, whole, ventral; G, dorsal portion of stylus removed, lateral; H, full stylus and part of penial sclerite removed; I, whole, ventrolateral oblique; J, stylus and muscle partially removed, dorsal; K, L, stylus partially removed, with K in ventrolateral oblique view, L in lateral view; M, N, coxites II and penial sclerite,partially dissected, with M in dorsal view, N in mesal (lateral) view.Abbreviations: CxtSty-art ¼ gonocoxite IIestylar articulation; Cxtpdp ¼ gonocoxite II posterodorsal process; CxtI ¼ "phallobase" (first gonocoxites); CxtII ¼ second gonocoxites (continuous with penis, forming "coxopenis" or CxtII þ Pen); Cxttvl ¼ gonocoxite II transverse lamella; Cxtvma ¼ gonocoxite I ventromedial apodeme; Pen ¼ penis and penial sclerite (continuous with CxtII, forming "coxopenis" or CxtII þ Pen); StIX ¼ sternum IX; Stap ¼ sternal apophysis; Sty ¼ gonostylus; Stydl ¼ gonostylar dorsal lobe; Stypdp ¼ gonostylar prosterodorsal process; Styvt ¼ gonostylar ventral tine; TgIX ¼ tergum IX; Tgap ¼ tergal apophysis. Muscle abbreviations indicated in Table 1.

Fig.16 in A general theory of genital homologies for the Hexapoda (Pancrustacea) derived from skeletomuscular correspondences, with emphasis on the Endopterygota

Fig.16. Genital skeletomusculature of Pycnopsyche antica (Trichoptera: Limnephilidae).A, genital and pregenital segments, lateral.B, genital and postgenital segments, posterior.C, same as B, left coxopod removed. D, same as C, lateral oblique. E, genital and postgenital segments, ventral. F, genital appendages, dorsolateral oblique. G, same as F, left gonopod removed. H, right gonopod and piece of postgenital complex, muscles removed, mesal (medial). IeK, penial sclerite and lateropenites: I, dorsolateral; J, dorsolateral, sheath membrane removed; K, ventrolateral, base of penial sclerite torn open. Abbreviations:?Ce ¼ structures possibly homologous with cerci; Cxa ¼ gonocoxa; Lpe ¼ lateropenite; Pen ¼ penis or penial sclerite; Prct ¼ proctiger; Seg ¼ segment; St ¼ sternum; Sty ¼ gonostylus; Tg ¼ tergum.Muscle abbreviations indicated in Table 1. Note: Segment numbers indicated with Roman numerals; addition signs indicate sclerite fusion.

Fig. 17 in A general theory of genital homologies for the Hexapoda (Pancrustacea) derived from skeletomuscular correspondences, with emphasis on the Endopterygota

Fig. 17. Key skeletomuscular morphological apomorphies of genitalia here inferred, mapped on the phylogenetic chronogram of the Hexapoda pruned to the specific terminals compared in the present study (tree modified from Misof et al., 2014). Bayesian analysis may provide estimates of rate and timing of inferred transformations.1. Hexapoda: male gonopore(s) situated on a lateromedially-undifferentiated gonopod (the penis). 2. Protura: male gonopores situated on distal (second) endopodal segment. 3. Collembola, Diplura: penes independently reduced to papillae. 4. Ectognatha: male and female genitalia derived from appendages (primary and secondary gonopods) of two successive segments. 5. Ectognatha: male phallic complex reduced to single, simple structure (the penis), modified from gonopores of abdominal segment X. 6. Dicondylia: dorsoventral penial extrinsic muscle lost. 7. Gonapophyses (endopods of coxopods IX) probably lost once in the Odonatoptera (along stem to Odonata) and once or twice in the Chiastomyaria. 8. Odonata: plesiomorphic condition of gamete transfer via spermatophore retained, although the latter is deposited in the autapomorphic secondary genitalia. 9. Ephemeroptera, Neoptera: sperm directly deposited in the female via intromission of the penis.10. Ephemeroptera: gonopods derive multiannulate styli.11. Ephemeroptera: penis laterally articulated with tergum IX, bearing derived tergopenial musculature. 12. Neoptera: sternopenial muscle duplicated, resulting in novel penial promotor. 13. Polyneoptera: secondary gonopods (coxopods IX) not incorporated into penial ("phallic") complex, being rather (usually) undifferentiated from sternum IX.14. Two independent lines of increasing complexity and disparity observed in the penial complex of Polyneoptera, once for the Zoraptera, the other for the Orthopterida (i.e., core Polyneoptera). 15. Two independent lines of penial complex simplification observed in Polyneoptera, once for the Plecoptera, the other for the Dictyoptera (namely, reduction observed in Isoptera).16. Grylloblattodea: coxopods IX secondarily differentiated from sternum IX.17. Eumetabola: gonostyli strengthened for clasping, bearing differentiated abductor and adductor muscles.18. Condylognatha: dorsal extrinsic penial muscle gained. 19. Hemiptera: gonopods undifferentiated from fused tergum and sternum IX, forming pygophore; pygophore also bearing origins of the penial extrinsic muscles. 20. Endopterygota: penis completely integrated with gonopods developmentally and gonopods, as in Hemiptera, bear the origins of the penial extrinsic muscles. 21. Endopterygota: penial musculature is duplicated dorsoventrally, resulting in greater functional potential of copulatory apparatus (penial-gonopodal complex). 22. Endopterygota: lateropenite ("parandrite" or "paramere" sensu Verhoeff) uniquely derived from penial sclerite (skeletomusculature of Psocodea in need of renewed study).23. Hymenoptera: origin of cupula (¼ "basal ring"), which controls overall motion of gonopods. 24. Hymenoptera: parossiculus derived from ventromedial gonocoxa, forming volsellar complex with lateropenite ("digitus"). 25. Sialida (sensu Bodreaux,1979 modified from Handlirsch,1908): gonocoxae migrated dorsally, fusing with tergum IX over the penis. 26. Sialida: external gonopore dissociated from the penial sclerites (probably in correlation with female loss of ovipositor). 27. Coleopterida: lateropenite lost. 28. Strepsiptera: genitalia reduced, with only a penis (or coxopenis) expressed. 29. Coleoptera: ancestral gonocoxa divided into two parts: the first or anterior gonocoxites ("phallobase") bearing the insertions of the extrinsic gonopodal musculature, the second or posterior gonocoxites incorporated to completely integrated with the penis (forming "coxopenis"); gonostyli variably associated with phallobase. 30. Euantliophora: genitalic complexity increases to extreme degree, extant taxa inheriting an autapomorphic aedeagal apodem (present in Siphonaptera and Nannochoristidae). 31. Siphonaptera: gonopodites, bearing styli, fused with tergo-proctiger complex. 32. Siphonaptera: sternum IX with lateral elongate arms. 33. Siphonaptera: sperm-pumping complex significantly modified. 34. Mecoptera: sperm-pumping complex with mittelplatte (note uncertainty of homology for boreid penial sclerites).35. Boreidae: sperm-pumping complex reduced, most slerites lost. 36. Boreidae: spermatophore regained. 37. Pistillifera: pistilltrö ager derived in sperm-pumping complex. 38. Diptera: partial to complete integration of sternum IX with gonopods; increasing genitalic complexity observed. 39. Amphiesmenoptera with relatively unmodified genitalia, although Lepidoptera with lateromedially fused lateropenites ("median plate") and enlarged and unmusculated gonostylus.

Fig.12 in A general theory of genital homologies for the Hexapoda (Pancrustacea) derived from skeletomuscular correspondences, with emphasis on the Endopterygota

Fig.12. Genital skeletomusculature of Apterobittacus apterus (Mecoptera:Bittacidae).AeC, genital and pregenital segments: A, lateral; B, distal; C, ventral.DeG, genital appendages and sternum IX: D, dorsal; E, lateral (partially dissected); F, lateral, left fragment of first coxite removed; G, same as F, with more muscle removed. H, left fragment of first coxite, mesal. IeN, complex of second coxite, lateropenite, and penial sclerite: I, lateral, first penial sclerite fragment removed; J, dorsal; K, dorsal, first penial sclerite fragment removed, dorsal.L, second coxites and first penial sclerite fragment removed,dorsal. M, same as L, dorsolateral oblique. N, first penial fragment,ventral (mesal), left and median tines partially broken. Abbreviations: Apd ¼ apodeme; Ce ¼ cercus; Cxa ¼ gonocoxa or first gonocoxites; Cxt# ¼ gonocoxite fragment; Lpe ¼ lateropenite; Pen# ¼ penial sclerite fragment; Prct ¼ proctiger; Sclbr-b-a'' ¼ sclerotic bridge between mittelplatte and second fragment of aedeagal apodem; Sclbr-cxt-cxt ¼ sclerotic bridge between first gonocoxites; SegVII ¼ seventh abdominal segment; Seg VIII ¼ abdominal segment VIII; StIX ¼ sternum IX; Sty ¼ stylus; TgIX ¼ tergum IX. Muscle abbreviations indicated in Table 1. Note: Sclerite fragment numbers indicated with Roman numerals; addition signs indicate sclerite fusion.

Fig.15 in A general theory of genital homologies for the Hexapoda (Pancrustacea) derived from skeletomuscular correspondences, with emphasis on the Endopterygota

Fig.15. Genital skeletomusculature of Tipula (Hesperotipula) californica (Diptera:Tipulidae). AeE, whole genitalia: A, lateral; B, dorsal; C, posteroventral oblique; D, distal; E, distal, tergum and sternum IX removed. F, tergum IX, mesal (ventral). GeI, genitalia with tergum IX and left gonopod removed, dorsolateral: G, with proctiger; H, proctiger partially removed; I, proctiger completely removed. JeL, right coxopod: J, K, mesal (medial); L, distal, oblique. M, genitalia with tergum IX and left coxopod removed, dorsolateral. N, right coxopod and penial sclerites, mesal. O, penial sclerites, lateral. Abbreviations: Ce ¼ cercus; Cxa ¼ gonocoxa; Lpe ¼ lateropenite; Pen ¼ penis; PenI ¼ first penial sclerite; PenI- nd ¼ penial needle; PenII ¼ second penial sclerite; PenIIdt ¼ dorsal tines of second penial sclerite; PenIImd ¼ medial disc of second penial sclerite; PenIII ¼ third penial sclerite; Prct ¼ proctiger; StVIII ¼ sternum VIII; StIX ¼ sternum IX; St-br ¼ sternal bridge; StCxa-sut ¼ sternocoxal suture; Sty ¼ gonostylus; Sty-dt ¼ dorsal tine of gonostylus; Sty- rcpt ¼ stylar receptor; Sty-vt ¼ ventral tine of gonostylus; TgIX ¼ tergum IX. Muscle abbreviations indicated in Table 1.

Fig. 3 in 21-Hydroxypregnane 21-O-malonylation, a crucial step in cardenolide biosynthesis, can be achieved by substrate-promiscuous BAHD-type phenolic glucoside malonyltransferases from Arabidopsis thaliana and homolog proteins from Digitalis lanata

Fig. 3. Size exclusion chromatography (SEC) and indication of relative 21MaT activity investigation pools III (A) and IV (B) of the ammonium sulfate precipitation.

Fig. 4 in 21-Hydroxypregnane 21-O-malonylation, a crucial step in cardenolide biosynthesis, can be achieved by substrate-promiscuous BAHD-type phenolic glucoside malonyltransferases from Arabidopsis thaliana and homolog proteins from Digitalis lanata

Fig. 4. Docking of modeled AtPMaT1 (ribbon diagram) with an overlay of the potential pregnane substrates (Sub) (shown in grey). The catalytic histidine (His) and the cosubstrate (CoS) are also shown.

Fig. 1 in 21-Hydroxypregnane 21-O-malonylation, a crucial step in cardenolide biosynthesis, can be achieved by substrate-promiscuous BAHD-type phenolic glucoside malonyltransferases from Arabidopsis thaliana and homolog proteins from Digitalis lanata

Fig. 1. Postulated biosynthetic pathway of cardenolide formation in Digitalis. The malonylation step [8] is marked by a rectangle. 1 Putative side chain cleaving enzyme (SCCE), 2 NAD:3β-hydroxysteroid dehydrogenase (3βHSD), 3 Δ4,5-3-ketosteroid-isomerase (3KSI), 4 progesterone-5β-reductase (P5βR), 5 NAD:3β-hydroxysteroid dehydrogenase (3βHSD), 6 putative pregnane 14β-hydroxylase, 7 putative pregnane 21β-hydroxylase, 8 malonyl coenzyme A:21- hydroxypregnane 21-O-malonyltransferase (21MaT).

Fig. 6 in 21-Hydroxypregnane 21-O-malonylation, a crucial step in cardenolide biosynthesis, can be achieved by substrate-promiscuous BAHD-type phenolic glucoside malonyltransferases from Arabidopsis thaliana and homolog proteins from Digitalis lanata

Fig. 6. Docking of homology modeled malonyltransferases with 3-O-acetylketol (displayed in grey) showing the distances between the catalytic histidine (His), the hydroxy group to be malonylated (Sub) and the malonyl residue presented by the co-substrate (CoA). A AtPMaT1 B AtPMaT2 C DlMaT1.

Fig. 5 in 21-Hydroxypregnane 21-O-malonylation, a crucial step in cardenolide biosynthesis, can be achieved by substrate-promiscuous BAHD-type phenolic glucoside malonyltransferases from Arabidopsis thaliana and homolog proteins from Digitalis lanata

Fig. 5. Expression of Dlmat1, Dlmat2, Dlmat3, and Dlmat4 in different plant tissues measured by real-time quantitative PCR (qPCR). Expression rates are standardized to the values of the actin transcript in each tissue and were displayed in relation to the expression in young leaves (set to equal 1) for each Dlmat gene.

opennotspecifiedSep 2023View details →

FIGURE 1 in Identification of ancient Cladocera-like fossils requires homologies: The Jurassic Kuqaia is not a Waterflea

FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with "posterior" appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (A–B and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E).

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