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Differences between generalized Paleozoic and modern pterygote nymphs. Fossils show the limb organ to be present on all body segments and serially homologous (=homonomous) in podites and rami (exites, endites). Up to 14 pairs of limbs occur; all bear climbing tarsi with once subsegmented ET and curved double claws. Abdominal ventral vesicles (formed from endites) are serially homologous with genitalia. All wings and winglets (=flattened epicoxal exites) are originally serial, fully articulated and mobile. Epicoxal pleuron is fused to cranium in the head, articulated to tergum and fragmented into wing pteralia in the thorax, and fused (with a suture) to terga in the abdomen (abdominal winglets are originally articulated, later fused, or reduced). In most modern nymphs the winglets plus their articulation were secondarily fused to terga and resemble lateral tergal extensions; limbs are much more dissimilar and appear to be not serially homologous. Schematic, after Kukalová-Peck (1991), updated. Kristensen (1998) and before interprets wings as autapomorphic in Pterygota and there is no appendage homologue in other Arthropoda; limb-derived appendages in mouthparts, thorax and abdomen are uniquely formed, not derived from a monophyletic all-arthropod limb ancestor; chewing lobes in mouthparts, vesicles, gonapophyses, penes and valvulae are of uncertain origin, not derived from all-arthropod endites and not serially homologous. If these were true, these profound morphological differences would remove insects from Arthropoda
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Most currently applied systematic methods use post-groundplan character states to reconstruct phylogenies in modern higher
Insecta/Arthropoda taxa. But, this approach is unable to separate synapomorphies from frequently occurring homoplasies. Conflicting,
unresolved and unrealistic higher-level phylogenies result. The reasons are analyzed. A contra...
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... homologous organs share a common genetic origin and similar construction, which is underlining all subse- quently added modifications. In the limb/wing organ system, these often include synapomorphies shared by the extant higher taxa (Figs. ...
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... single piece (not a coxopodite); the so-called endites in the head as secondary, non-serial lobes; the maxilla as not more primitive than the thoracic leg. But, these are errors since sutures between podites left behind after their fusion are retained in mandibular coxopodites of some trilobites, and are quite distinctive in living Archaeognatha (Figs. 11, 12); the thoracic leg, Z-shaped, suspended from two flattened pleura and bearing a wing as well as climbing tarsus, is adapted to lift the insect body up and to serve equally well in walking, running over uneven substrates, climbing, jumping, holding, scratching, and flying, and is a marvel of multiple functionality even among arthropod ...
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... shifted upward into coxa (CX), enclosed, immobilized, and used as tactile appendage (three autapomorphies of Archaeognatha). After Kukalová-Peck (1998), altered. Kristensen (1998 and before maintained that coxal lobe is not an exite because it is positioned in the middle of coxa. This is an error because some arthropod exites shift upward (as in Fig. 12 here), from the membrane into the proximal podite, into which they eventually become enclosed (E. L. Smith, communication during cooperation with JKP) have strongly regressive influence in higher taxa (suborders and upward taxa). Note, that to save space, all currently used systematic methods limited to exploring higher-level ...
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... (see Fig. 17.1). Never used in aerial forward flapping flight, the prothoracic winglets retained the original venation of the pre-flight ancestral protowings, which originally may have formed a continuous homonomous series on all segments of the thorax and abdomen. After JKP in: Wootton and Kukalová-Peck (2000) absorbing moisture (Figs. 2, 15), and in genitalia as gonapophyses, penes, and ovipositor valves for copulating nad laying eggs (Figs. 2, 4, 15, 20, 21). Coxal endites articulate coxae to the thoracic sternum in ...
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... the original venation of the pre-flight ancestral protowings, which originally may have formed a continuous homonomous series on all segments of the thorax and abdomen. After JKP in: Wootton and Kukalová-Peck (2000) absorbing moisture (Figs. 2, 15), and in genitalia as gonapophyses, penes, and ovipositor valves for copulating nad laying eggs (Figs. 2, 4, 15, 20, 21). Coxal endites articulate coxae to the thoracic sternum in ...
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... pleural membranes were found by us to be supported by flattened podites. Three separate pleura (subcoxal, coxal, and trochanteral) were distinctly delim- ited by sutures in the abdomens of fossil Diaphanopterodea (Fig. 20). These, and abdominal structures in modern Ephemeroptera and extinct and extant Zygentoma, were instrumental in reconstructing the ancestor (groundplan) of the abdomen in Dicondylia (Fig. 15), which also shows the position of abdominal endites as in Fig. 1. In Ptery- gota, Paleoptera (Palaeodictyopterida + (Odonatoptera + ...
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... loss and ''disappearance'' of plesiomorphic character states are a common evolutionary transformation. The adage ''evolution is reduction and tin- kering with the rest'' is widely considered valid. In arthropod appendages, there is enormously varied reduc- tion/fusion in the number of limb podites, rami and wing veins in the modern entomofauna (Figs. 1-21). Trilobites were also much more complex in the Cambrian than they were in the Permian (Webster 2007). Countless other examples exist. Post-groundplan reductions, fusions, parallelism and homoplasies especially target and confuse the serial homology of limb-derived appendages, and superficially defy their provenance from a single all- ...
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... engineering and ''supported'' three phylogenies: (Paleoptera + Neop- tera); (Ephemeroptera + (Odonatoptera + Neoptera)); and (Odonatoptera + (Ephemeroptera + Neoptera)) (Kukalová-Peck 1983; see below). In contrast, clues from fossils showed that ancestral sclerites were quite small, numerous, and arranged in regular rows protecting blood channels (Figs. 2, 14, 18.). In Neoptera, they were assembled into three irregular, oblique clusters (the first, second, and third axillary), while in Paleoptera all fusions followed an arrangement into rows (Figs. 14, 17-19). Axillaria and articular plates in some modern insects still bear weak sutures delimiting the original, small sclerites. But, without ...
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... , 1998Beutel and Pohl 2006). The problem is that arthropod evo- lutionary morphology is still little known. Across all arthropod orders, so far, only the insect wing and most of the limb structure have been fully homologized (but the limb derived insect hypopharynx and some derived insect geni- talia are still inadequately understood) (Figs. 1-21) (Kukalová-Peck 1991Haas and Kukalová-Peck 2001;Kukalová-Peck and Lawrence ...
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... a number of features, which were previously unknown or misinter- preted: the presence of the epicoxal pleuron shared by all Arthropoda; the identity of insect clypeolabrum; the composition of mandible; maxilla; and labium in the head (Fig. 6), subcoxal pleuron in the thorax (Figs. 7-9), and subcoxal + coxal + trochanteral pleuron in the abdomen (Figs. 15, 20). In the thorax, epicoxal pleuron and its exite were identified as the homologue of the wing articulation and flattened wing ramus (Figs. 7-9). In the abdomen, it showed serial pregenital leglets (starting at prefemur (PFE) and primitively bearing double claws) as homologues of the previously enigmatic abdominal ''styli'', vesicles and ...
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... flattened wing ramus (Figs. 7-9). In the abdomen, it showed serial pregenital leglets (starting at prefemur (PFE) and primitively bearing double claws) as homologues of the previously enigmatic abdominal ''styli'', vesicles and gonostyli; gonocoxites in male and female genitalia were recognized as coxopodites, and genital appendages as endites (Figs. 15, 19, 20) (Kukalová-Peck and Richardson 1983;Kukalová-Peck 1983, 1985, 1987Riek and Kukalová-Peck 1984;Haas and Kukalová-Peck 2001;Kukalová-Peck and Lawrence ...
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... systematic rule determining that inside every higher pterygote taxon, significant veinal fusions and reductions can be only added and never removed are documented here in Figs. 1-21, and in many publications ( Haas and KukalováPeck 2001;Kukalová-Peck 1969, 1978, 1983, 1985, 1987Brauckmann 1990, 1992;Kukalová-Peck and Lawrence 2004;KukalováPeck and Richardson 1983;Riek and Kukalová-Peck 1984;Sharov 1957Sharov , 1966Smith 1970Smith , 1988. The irreversibility rule plays an important role in higher-level phylogenies. ...
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... easily tested ground- plan characters, such as that all blattoids have anojugal lobe in the hind wing starting at the anal fold, while in orthop- teroids and plecopteroids it starts always at the claval fold; all Neoptera have three identically constructed axillary sclerites, etc. Examples of the irreversibility of groundplans are countless (see Figs. 1-21). Raff (1996), Marshall et al. (1994) and Raff (1996) have analyzed genetic attributes and evolutionary constraints versus irreversibility of the old Paleozoic ...
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... Paleozoic-based features most frequently ignored by the advocates of post-groundplan methods are exites, endites, sutures denoting fusions between ''disappeared'' podites, abdominal pleural plates, abdominal vesicles, telopodite-like palps, styli and gonostyli, and any other feature showing serial homology of the limb-derived appendages (compare Figs. 1-20). These critics often cite the experiments in Drosophila, suppression and desup- pression, and claim that they ''disprove'' the irrreversibility of ordinal character states. But, geneticists have other explanations for these phenomena and give their support for irreversibility of the old groundplans ( Raff et al. 1991;Marshall et al. ...
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... function. Example: The arch-conservative order Archaeognatha (jumping bristle- tails) has a cryptic life style and lives in crevasses and litter. From the Devonian to modern times, it has kept its unreduced, telopodite-like palps, which are used in climbing and for balance (Fig. 10.1), articulated epipleuron (Figs. 8, 9), up to three exites (Figs. 8-12) and other Paleozoic character states. Some Hymenoptera (sawflies, Fig. 5) use plesiomorphic multi-segmented palps for raking pollen and retained not only unreduced, Hexapoda-level maxillary telopodites, but also curved double claws (which some Paleozoic insects bore on up to 15 pairs of limb appendages, Fig. 13). Before the Great ...
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... of the post-groundplan method in inter- preting higher-level phylogenies. Those authors explicitly reject two decades in limb/wing contributions: all-arthro- pod ancestral polyramous limb (Fig. 1), the seriality of mouthparts and endites (Fig. 6), mandible and maxilla formed from a three-segmented coxopodite (rather than from ''coxa'', but see Figs. 6, 11, 12), origin of genital appendages and vesicles from endites (see Figs. 15, 20, 21), homology of wings and the limb rami in Archaeog- natha from exites (see Figs. 2, 10, 12), and of mandibles and maxillae in Hexapoda from coxopodites (see Fig. 6), etc. Instead, they see all exites and endites as occasionally developing secondary lobes, ...
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... polyramous limb (Fig. 1), the seriality of mouthparts and endites (Fig. 6), mandible and maxilla formed from a three-segmented coxopodite (rather than from ''coxa'', but see Figs. 6, 11, 12), origin of genital appendages and vesicles from endites (see Figs. 15, 20, 21), homology of wings and the limb rami in Archaeog- natha from exites (see Figs. 2, 10, 12), and of mandibles and maxillae in Hexapoda from coxopodites (see Fig. 6), etc. Instead, they see all exites and endites as occasionally developing secondary lobes, mandibles and maxilla as a single segment (coxa), etc., exactly as Manton (1977) in ''Uniramia''. However, these antiquitated, long rejected- but lately resurrected ...
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... In all Animalia, ancestral characters were reduced in most, but often still occur in some modern species, the same way as in Insecta, Crustacea and Chelicerata. In evolutionary morphology, their published un-reduced occurrences are considered the definitive proof of their existence, and a verifiable plesiomorphy which must not be ignored (see Figs. 5, 6, 8-10, 12, 19-21, and text). The equally impor- tant complementary evidence in modern ontogeny, embryology, and genetics published by various authors (see references cited by Kukalová-Peck 1978, 1983and Kukalová-Peck and Lawrence 2004) also must not be ignored, because it provides all-important cross-checking and additional invaluable information. As a fact, a ...
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... plesiomorphies found in modern insects (Figs. 2-21, text), homologous structures in other arthro- pods, and compatible data from other biological fields, embryology, ontogeny, genetics, developmental genetics, and experiments with transplants, provide the necessary verification of the oldest, often-unfamiliar fossil character states (see examples below). Using many sources is also highly ...
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... small, equal articles for flexibility in climbing, which disguised two podites, basitarsus and (subdivided) eutarsus. Thus impoverished and transformed, grasshopper leg shows only a distant resemblance to the much better preserved maxilla, and none to the mandible, flagellated antennae and cerci, and reduced abdominal styli and gonostyli (Figs. 2, 6, 15, 20). The Snodgrass model also shows no clues to either the origin of ''teeth'' in the mouthparts, working parts of genitalia, abdominal plate gills, vesicles or styli, or of the wings (Figs. 1-6, 10, 11), and it show all these appendages as ''secondary'' (and, quite useless for higher classification!!). It is because of this concept, which ...
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... limb model, and interpreted the arthropod mandible incorrectly as a ''single'' coxal podite (but, it is composed of three podites: Fig. 6). After that she correctly noticed that the most primitive modern insects (Archaeognatha: jumping bristletails) bear mandibles dis- sected by sutures marking their composition from several limb podites (Figs. 11.1, 12.1). This incorrectly convinced her that unlike Arthropoda, the insects eat with the ''tip of their legs'' in the same way as Onychophora. Conse- quently, Manton disassembled the monophyletic phylum Arthropoda, and referred Atelocerata (=Myriapoda + Hexapoda/Insecta) plus Onychophora to a new (but poly- phyletic) ''uniramous'' phylum ...
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... to a new (but poly- phyletic) ''uniramous'' phylum ''Uniramia''. In reality, all arthropods are polyramous and eat with the limb base (coxopodite), which is composed of three limb podites and two endites (Figs. 1, 6); however, only Archaeognatha retained the sutures that separate these three podites (plus a coxal subdivision) and two endites (Figs. 11.1, 12.1). Errors coming from a well-known scientist can have a long half-life. It took 17 years to rectify this massive miscon- ception (Kukalová-Peck 1983Wägele 1993). Now, by ignoring the need for a monophyletic all- arthropod limb ancestor and multiple evidence for its serial expression, the post-groundplan method is again bringing back the ...
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... back the problem with limb homology, in the contributions of Kristensen (1995Kristensen ( , 1998, Willmann (1997), andGrimaldi andEngel (2005) (see examples and discussions below). In the book of the last authors, the limbs in Hex- apoda are once again defined as ''uniramous'' and in Crustacea as ''biramous'', rather than both being polyra- mous (Figs. 8, 12.1-4, 15). All rami-based appendages are ''secondary lobes'', and the concept of serial homology of limbs, repeatedly confirmed in paleontology, develop- mental genetics, and classical genetics, is abandoned to a great loss of synapomorphies shared by the higher arthropod taxa. But, the post-groundplan approach, with its emphasis on numbers, ...
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... the post-groundplan approach, this fundamental split of Pterygota was rejected based on an erroneous and out- dated belief that mandible and maxilla are composed of a single podite ''coxa'', bearing two ''secondary'' lobes (mola and incisor in mandible, lacinia and galea in maxilla) (but, see Figs. 6, 8.1, 11, 12). This faulty model has many erroneous repercussions; among others it suggests a dif- ferent phylogeny when applied to the mandible, than when applied to a maxilla. Kristensen (1991Kristensen ( , 1995 compared the mandible of Odonatoptera and Neoptera and found the following five ''synapomorphies'': massive, broad mandible; tight ball ...
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... in fossils: Flexible, once subsegmented climbing tarsus plus double-clawed pretar- sus occurs on up to 15 limb pairs of Paleozoic insects: in two pairs of palps, three pairs of thoracic legs, eight pairs of abdominal leglets, one pair of gonostyli, and one pair of cercopods (the latter, only in Cercopoda, Fig. 13) (Kukalová-Peck 1983, 1987 (Figs. 1-21). Evidence in extant insects: Hexapod climbing tarsus without claws is retained in the maxillary palps of modern Archaeognatha (Figs. 7, 8); with claws, it occurs in Hymenoptera, in the sawfly Pleroneura sp. (Fig. 5; the fused-immobile claws in palps rake pollen), and in the ''walking pupa'' of Raphid- ioptera (E. L. Smith, personal ...
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... eutarsal subdivisions and their fluctuations are limited only to the thoracic legs involved in climbing and walking (Figs. 2-5). Outside the thorax, tarsi in the stem- species that survived the Great end-Permian Extinction are almost always reduced, many times independently and in parallel. Many Paleozoic insects show the presence of 3 tarsal joints ...
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... groundplan method (applied to the limb/wing organ system, see examples in Figs. 1-21) uses only one ancestor and one set of irreversibility rules for all higher taxa in Arthropoda. Therefore, it can produce only one higher phylogeny (right or wrong). For winged insects it ...
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... offers the following evidence: Paleontology: In the plesiomorphic Paleozoic juveniles of Paleoptera and Neoptera, the winglets are fully mobile and articulated in a functional (lateral) position, as in miniature adults (Figs. 2, 14). Several thousands of detached nymphal winglets of Ephemeroptera from the Early Permian from Elmo, Kan- sas, are deposited at the Museum of Comparative Zoology, Harvard University, Cambridge (modern nymphal wing pads stay always fused to terga). About 200 Paleozoic juveniles with fully articulated, mobile wings are recorded (Fig. 14). ...
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... wings are homologous with the outer rami (=ex- ites) of arthropod epicoxal pleura (Fig. 17.1-2). The flattened protowing exite travelled dorsally (upward) on its epicoxal pleuron, which fused completely underneath it and above the ventral wing process (the second axillary is doubled, Fig. 17). Note that a similar upward shift and fusion occurs in the coxal exite of modern Archaeognatha and other arthropods. This evaginated in the ...
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... ta) (Kukalová-Peck 1987, Figs. 15-17;) also entered the same habitat and diverged from the free-living main stock, but only after their protowings were larger and more advanced. Therefore, the latter became fused to the terga as wing-based sidelobes separated by deep sutures (similar as prothoracic sidelobes and wing pads in modern nymphs (Fig. 2), but sutures are retained only in modern nymphal peloridiids). Evidence: Structure: Modern silver- fish bear inside their thoracic sidelobes the wing tracheae with a veinal branching pattern, which shows them as derived from wings (S ˇ ulc 1927); in Paleozoic silverfish, they are separated from terga by deep sutures like in pro- ...
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... only with fully homologized arthropod characters. Besides wings, so far only the limb organ sys- tem ( Fig. 1) is ready to produce independently reconstructed phylogenies. In this, Beutel and Pohl (2006) overlooked the now available data: for instance confirma- tion of the wing-based basal split in Neoptera in the divergence of female genitalia (Fig. 20, here). As shown by Sharov (1957Sharov ( , 1966 and Hennig (1981), the ovipositor third valve (gonoplac) in Neoptera is formed in two dif- ferent ways: either by the stiffened gonostyli (in Orthoneoptera + ancestral Pleconeoptera), or by the elongated gonocoxites (in Blattoneoptera + Hemineopter- a + Endoneoptera). The authors also overlooked ...
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... estab- lished, a comparison of groundplan character states between analyzed orders (or other higher taxa) usually instantly reveals the synapomorphies shared by ordinal sister groups. The groundplan method was applied to the arthopod limb/wing organ, which was gradually upgraded for the last 30 years and is offered here in several evolu- tionary models (Figs. 1, 15, 17-20). Its potential in Insecta is still far from being exhausted, and it has not yet been broadly applied to Crustacea and Chelicerata in published contributions. ...
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... of (Neuropterida + Mecopterida), and Neoptera as split into two super-lineages: (Pleconeoptera + Orthoneoptera) and (Blattoneoptera + (Hemineoptera + Endoneoptera)). Both relationships were reinforced by other character sets, shared irreversible synapomorphies in veinal fusions and braces, and two different origins of the third ovipositor valvula (Fig. ...
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... taxa: Mandibulata, Atelocerata, Parain- secta, Insecta. In Pterygota, the most debated phylogenies concern pterygote basal divisions, lineages and basal orders in these lineages. It is proposed in this account that these relationships are all solvable by the groundplan method in an objective and repeatable way (see documentation above and in Figs. ...
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... same old, ''direct'', but naive approach to limb evolution and homology exploded Arthropoda and started a massive debate unparalleled in the history of modern biology. As during the heady era of numerical taxonomy, the lure of numbers is again eroding an evolutionary approach, in spite of strong and ever growing evidence supporting evolution (see Figs. 1-21 ...
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A short article describing the gall-inducing and leaf-mining fauna of the nature reserve Strødam, north of Copenhagen, Denmark. In Danish.
Citations
... Another indicator can be discerned through the presence of exites and endites, which are pleiomorphic traits among all crustaceans (J. Kukalova-Peck et al. 2007). Detecting these exites and endites assimilated into the body assists in identifying absent components. ...
... The presumed ancestral xenocarid likely possessed appendages of the same type as cephalocarida ( Figure 6 and Figure 31), featuring the characteristic 3-podomere exopodite and complete podomeres reminiscent of the basal crustaceans (D. Walossek et al. 1993;J. Kukalova-Peck et al. 2007). The posterior end of the body culminated in two furca, terminating with flagella. This hypothetical precursor gave rise to two species that occupy different niches, cephalocarida (Figure 23 and Figure 24) and the common ancestor of remipedia ( Figure 21 and Figure 22) and SDSNH 28852 (Figure 19 and Figure 20). Cephalocarida evolved wi ...
Through a comprehensive investigation of SDSNH 28852, previously found inside xenocarida, this work elucidates the evolutionary relationship between xenocarida and hexapoda. Reexamining M. J. Emerson's and F. R. Schram's original data reveals inconsistencies in their descriptions. Molecular data and physical characteristics clearly link xenocarida to hexapoda. SDSNH 28852 plays a key role, sharing ancestors with remipedia and showing significant similarity with cercopoda. Adaptations that aid in the terrestralization of hexapoda, emphasizing breathing processes and appendage transformations into specialized feeding tools. This study suggests important research directions. Mutations in remipedia might be validated by genetic research, and micro-CT scans of SDSNH 28852 would provide a 3D image of appendage structures. SDSNH 28852 remains a significant resource under the care of Dr. Michael S. Engel. By improving previous data, this work improves the xenocarida-hexapoda relationship. It contributes to evolutionary biology by offering information on the emergence and adaptation of terrestrial arthropods. The complex relationship between xenocarida and hexapoda arises, describing adaptation and diversification as a result of environmental change.
... Accordingly, finding successive series of different instars attributable to single species is rare among Paleozoic fossils (e.g., 143,158). Hypotheses regarding the postembryonic development of Paleozoic hexapods and particularly of winged insects (Pterygota) were greatly influenced during the past 40 years by the studies of Kukalová-Peck (69,70,73,75,76), which were partially based on earlier works by Sharov (148,149). These were later either adopted without critical revision of examined specimens or partly disputed on the grounds of unconvincing evidence (10,15,47,65,126,139,146). Given this state of affairs, our view of postembryonic development in Paleozoic hexapods relative to that of extant ...
While Mesozoic, Paleogene, and Neogene insect faunas greatly resemble the modern one, the Paleozoic fauna provides unique insights into key innovations in insect evolution, such as the origin of wings and modifications of postembryonic development including holometaboly. Deep-divergence estimates suggest that the majority of contemporary insect orders originated in the Late Paleozoic, but these estimates reflect divergences between stem groups of each lineage rather than the later appearance of the crown groups. The fossil record shows the initial radiations of the extant hyperdiverse clades during the Early Permian, as well as the specialized fauna present before the End Permian mass extinction. This review summarizes the recent discoveries related to the documented diversity of Paleozoic hexapods, as well as current knowledge about what has actually been verified from fossil evidence as it relates to postembryonic development and the morphology of different body parts.
... The preoral segment is marked by vax, nk2-1, rx, dlx, bf-1, and otp expression (in vertebrates, these markers are expressed in the premandibular segment and forebrain anlage). The oral (tentacle) segment is marked by otx, pax6, emx, barH, dbx, irx, lim 1/5, and en expression; in Lophotrochozoa, these markers are expressed in the tentacle (peristomial) segment, and in deuterostomes, in the collar segment (in vertebrates, these markers are expressed in the mandibular somite, the spiracle segment and rudiment of the midbrain), and in arthropods, in the deutocerebral segment (Finkelstein and Boncinelli, 1994;Acampora et al., 1995Acampora et al., , 1998Acampora and Simeone, 1999;Arendt and Nübler-Jung, 1999;Seo et al., 1999;Murakami et al., 2001Murakami et al., , 2002Hirth et al., 2003;Lowe et al., 2003Lowe et al., , 2015Kuratani, 2004Kuratani, , 2005Kuratani et al., 2005;Arenas-Mena, 2006, 2008 Gąsiorowski and Hejnol, 2020;Andrikou and Hejnol, 2021). Along with the morphological data, this information emphasizes the homology of the tentacle (oral) segment and the associated tentacle apparatus wihin all taxa of triploblastic Bilateria (Fig. 17). ...
... Another point of view is that the wings are somehow connected with pleural appendages of the limbs such as Ephemeroptera tracheal gills or Apterygota styles (Oken, 1831;Gegenbaur, 1874;Osborn, 1905;Woodworth, 1906;Wigglesworth, 1973Wigglesworth, , 1976. The assumed origin of the wings from pleural appendages of the limbs was supported in the studies on the Paleozoic insect morphology (Kukalová-Peck, 1985, 1991, 1992, 2008Kukalová-Peck and Lawrence, 2004;Kukalová-Peck et al., 2009). In the insects, imaginal discs of the limb and wing (and haltere) are formed as a single rudiment with the ventrolateral position, then they are separated, and the wing disc (and haltere) shifts to the dorsal side (Tower, 1903;Auerbach, 1936;Goto and Hayashi, 1997). ...
... The most important development in the understanding of the origin and evolution of insect wings is to be found in the works of the paleoentomologist Jarmila Kukalová-Peck (1978, 1983, 1991, 2008 and summarised in Haas & Kukalová-Peck (2001) and Kukalová-Peck & Lawrence (2004). This system is based on the recognition of a protowing groundplan (Fig. 1A) based on a detailed study of pterothorax, pteralia and wings of numerous extant and extinct insects and a hypothesis of wing origin from "limbderived, locomotory, plate-like appendages, such as the crustacean swimming uropods". ...
The hind wings of all known families and most subfamilies of Coleoptera are illustrated, annotated and discussed utilising the terminology of Kukalová-Peck and Lawrence (2004), with a few changes in nomenclature suggested by the senior author. The beetle families are discussed in 21 groups, based on recent classifications of Coleoptera. For each of these groups, the most recent works on phylogeny and classification are reviewed, and the wing characters are discussed to determine if some of the wing features might support or refute relationships based on recent molecular and morphological analyses. Part 1 includes a general discussion of wing structure divided into the following sections: hind wing fields, veinal systems (including the history of wing nomenclature), wing folding, wing edge and embayments, hinges and bending zones, cross-veins and braces, cells and other landmarks. It is followed by discussion of the first 14 groups (Archostemata to Elateroidea), 15 figures supporting general discussions, and 426 labelled wing images of the discussed groups, representing 380 genera.
... Nonetheless, this argument is also not strong enough because the second character state is also present in some genera of Nymphidae and the third character state is absent in most antlions. Given the complex and sometimes unpredictable evolutionary pattern of insect wing venations 23 , the presently used wing characters may not provide sufficient phylogenetic signal to resolve the higher phylogeny of Myrmeleontoidea, especially including many fossil taxa. However, the morphological arguments in Makarkin et al. 7 placing Babinskaiidae together with Nymphidae in Nymphidoidae, i.e., the presence of trichosors and the completely separated forewing MP and CuA, are obviously attributed to the plesiomorphic condition in Neuroptera, which was also mentioned by these authors. ...
Babinskaiidae is an extinct family of the lacewing superfamily Myrmeleontoidea, currently only recorded from the Cretaceous. The phylogenetic position of this family is elusive, with inconsistent inferences in previous studies. Here we report on three new genera and species of Babinskaiidae from the mid-Cretaceous Kachin amber of Myanmar, namely Calobabinskaia xiai gen. et sp. nov., Stenobabinskaia punctata gen. et sp. nov., and Xiaobabinskaia lepidotricha gen. et sp. nov. These new babinskaiids are featured by having specialized characters, such as the rich number of presectoral crossveins and the presence of scaly setae on forewing costal vein, which have not yet been found in this family. The exquisite preservation of the Kachin amber babinskaiids facilitate a reappraisal of the phylogenetic placement of this family based on adult morphological characters. Our result from the phylogenetic inference combining the data from fossil and extant myrmeleontoids recovered a monophyletic clade composed of Babinskaiidae and another extinct family Cratosmylidae, and further assigned this clade to be sister group to a clade including Nemopteridae, Palaeoleontidae, and Myrmeleontidae. Babinskaiidae appears to be a transitional lineage between Nymphidae and advanced myrmeleontoids, with ancient morphological diversification.
... As one of the most enigmatic evolutionary innovations, wings enabled insects to take to the sky and radiate into previously inaccessible niches. Insects typically possess two pairs of wings located on the second and third thoracic segments (T2 and T3), connected to the body wall via a complex hinge mechanism (fossil Palaeodictyoptera also possessed superficially wing-like structures on other segments; however, these winglets lacked hinges and are not considered true wings [5]; but see [6]). Thus, bona fide wings are restricted to T2 and T3, and their ancestral affinities to other structures did not reveal themselves until more recent work succeeded in reconciling two competing hypothesis on the evolutionary origin of wings. ...
Modification of serially homologous structures is a common avenue towards functional innovation in developmental evolution, yet ancestral affinities among serial homologues may be obscured as structure-specific modifications accumulate over time. We sought to assess the degree of homology to wings of three types of body wall projections commonly observed in scarab beetles: (i) the dorsomedial support structures found on the second and third thoracic segments of pupae, (ii) the abdominal support structures found bilaterally in most abdominal segments of pupae, and (iii) the prothoracic horns which depending on species and sex may be restricted to pupae or also found in adults. We functionally investigated 14 genes within, as well as two genes outside, the canonical wing gene regulatory network to compare and contrast their role in the formation of each of the three presumed wing serial homologues. We found 11 of 14 wing genes to be functionally required for the proper formation of lateral and dorsal support structures, respectively, and nine for the formation of prothoracic horns. At the same time, we document multiple instances of divergence in gene function across our focal structures. Collectively, our results support the hypothesis that dorsal and lateral support structures as well as prothoracic horns share a developmental origin with insect wings. Our findings suggest that the morphological and underlying gene regulatory diversification of wing serial homologues across species, life stages and segments has contributed significantly to the extraordinary diversity of arthropod appendages and outgrowths.
... Formicidae mandible microstructure likely mirrors this concept. Furthermore, from a palaeontological and evolutionary perspective, it might also be that an effective Bauplan for ant mandible construction arose in the Cretaceous (Nel, Perrault, & Néraudeau, 2004) or was retained from the ancestral hexapod stock that gave rise to modern Formicidae (see discussion in Kukalová-Peck, 2008). Ant mandible microstructure therefore potentially represents an example of stasis and conserved evolution at the microscopic scale (Eldredge et al., 2005). ...
Exoskeletons characterise Arthropoda and have allowed the morphological and taxonomic diversity of the phylum. Exoskeletal sclerotisation occurs in genetically designated regions, and mandibles represent one such area of high sclerotisation. Mandible morphology reflects dietary preferences and niche partitioning and has therefore been well documented. However, mandibular cuticular microstructure has been under‐documented. Here we use scanning electron microscopy to explore mandible microstructure in four disparate Australian Formicidae taxa (ants) with different life modes and diets: Camponotus nigriceps, Iridomyrmex purpureus, Odontomachus simillimus and Rhytidoponera aciculata. We test the hypothesis that mandible construction is highly conserved across these species, as would be expected for arthropod cuticle. We show broadly similar mandible microstructure but report that pore canals and cuticular indentations are not ubiquitous among all studied taxa. Our preliminary results demonstrate that ant taxa have morphologically plastic mandibles with a highly conserved construction, potentially reflecting an interesting record of evolutionary stasis.
... However this structure is subdivided internally into three sub-basivenales by weaker membranes (Fig. 2a), one for MA (MBa), one for MP (MBp) and two for Cu (CuBa and CuBp) (Suppl. movies [12][13][14][15][16][17]; the MA's and the MP's are basally fused together into a MB. MA is clearly going into the same vein as R, but their basivenales are independent; the composite vein RA + RP + MA contains the three veins separately in its basal part near the transverse vein Ax0, but these three veins are distally fused into a unique vein, before RP + MA separates again into the arculus. ...
Being implied in flight, mimetism, communication, and protection, the insect wings were crucial organs
for the mega diversification of this clade. Despite several attempts, the problem of wing evolution
remains unresolved because the basal parts of the veins essential for vein identification are hidden in
the basivenal sclerites. The homologies between wing characters thus cannot be accurately verified,
while they are of primary importance to solve long-standing problems, such as the monophyly of the
Palaeoptera, viz. Odonatoptera, Panephemeroptera, and Palaeozoic Palaeodictyopterida mainly
known by their wings. Hitherto the tools to homologize venation were suffering several cases of
exceptions, rendering them unreliable. Here we reconstruct the odonatopteran venation using fossils
and a new 3D imaging tool, resulting congruent with the concept of Riek and Kukalová-Peck, with
important novelties, viz. median anterior vein fused to radius and radius posterior nearly as convex
as radius anterior (putative synapomorphies of Odonatoptera); subcostal anterior (ScA) fused to
costal vein and most basal primary antenodal crossvein being a modified posterior branch of ScA
(putative synapomorphies of Palaeoptera). These findings may reveal critical for future analyses of the
relationships between fossil and extant Palaeoptera, helping to solve the evolutionary history of the
insects as a whole.
... In this work, I follow the wing venation nomenclature of Kukalová-Peck (1983), amended by Kukalová-Peck (1991, 2008, also contributions by Riek and Kukalová-Peck (1984), Nel et al. (1993), Bechly (1996), and Petrulevičius and Gutiérrez (2016). The higher classification of fossil and extant Odonata is based on the phylogenetic system of Bechly (1996Bechly ( , 2007. ...
A new burmagomphid anisopteran, Satelitala soberana gen.
et sp. nov. is described from the lower Eocene of Laguna del
Hunco, Patagonia, Argentina. The new genus is characterised
by hindwing characters such as the subdiscoidal triangle
not elongated; anal loop divided longitudinally; paranal
cell divided longitudinally; five terminal cells between RP
and MA; five terminal cells between MP and CuA; and
obtuse angle between PsA and CuP+AA. Burmagomphid
dragonflies were represented so far only by one specimen
from the middle Cretaceous of Southeast Asia. This new record
extends the distribution to Patagonia, to the Cenozoic,
and also to paleolake deposits.
... The origin, development and dynamics of an intriguing subject in biology, insect winged flight, is one of the most exciting topics in organismic biology [1][2][3][4][5][6][7][8][9][10]. Some authors have suggested that the first winged insects had living tissue inside wing-like specialized structures and used to skim or row on the surface of water [9,10]. ...
Some consider that the first winged insects had living tissue inside the wing membrane, resembling larval gills or developing wing pads. However, throughout the developmental process of the wing membrane of modern insects, cells and tracheoles in the lumen between dorsal and ventral cuticle disappear and both cuticles become fused. This process results in the rather thin rigid stable structure of the membrane. The herewith described remarkable case of the dragonfly Zenithoptera lanei shows that in some highly specialized wings, the membrane can still be supplemented by tracheae. Such a characteristic of the wing membrane presumably represents a strong specialization for the synthesis of melanin-filled nanolayers of the cuticle, nanospheres inside the wing membrane and complex arrangement of wax crystals on the membrane surface, all responsible for unique structural coloration. © 2017 The Author(s) Published by the Royal Society. All rights reserved.
