Content uploaded by Jarmila Kukalová-Peck
Author content
All content in this area was uploaded by Jarmila Kukalová-Peck on Dec 17, 2014
Content may be subject to copyright.
A preview of the PDF is not available
New Carboniferous Diplura, Monura, and Thysanura, the hexapod ground plan, and the role of thoracic side lobes in the origin of wings (Insecta)
Abstract
A new Upper Carboniferous japygid, Testajapyx thomasi n.gen. et n.sp., shows that only Diplura of Entognatha shared an ancestral ground plan with Insecta–Ectognatha. Pleuron, palps, thoracic and abdominal legs, and vesicles are compared between Diplura, Archeognatha, Monura, Thysanura, and Pterygota. The origin and evolution of side lobes on the head, thorax, and abdomen, and of cereal legs, cerci, and posttarsal (=pretarsal) claws are considered. The paleoenvironment of the earliest insects and its impact on evolution of thoracic side lobes is discussed. Consistent evidence from several biological disciplines shows that protowings evolved from thoracic side lobes, which probably originated from serial, articulated outer appendages (=exites) of the arthropodan leg. Exites were continuously active and functional and became preadapted for flapping movements. Flight developed as the pterygotes diversified. Basic wing structures are monophyletic. Flight adaptation was finished and differences in wings arose somewhat later, by various means and in parallel, in already diversified lineages of Pterygota.
... Diverse spore-and pollen-rich coprolites suggest that palynivory was well established in equatorial peat swamp ecosystems during the Middle-Late Pennsylvanian. The coprolite record of palynivory is supplemented by rare occurrences of insect body fossils with gut contents from Mazon Creek in the Illinois Basin (Middle Pennsylvanian, Moscovian) Kukalová-Peck 1987;Shear and Kukalová-Peck 1990). These specimens provide direct evidence of lycopsid spore consumption across multiple insect lineages, including Eucaenus ovalis Scudder (Archaeorthoptera: Eucaenidae) with probable Lycospora spores (Carpenter and Richardson 1976;Scott and Taylor 1983), an unidentified archaeorthopteran with Cappasporites distortus Urban (Lepidodendrales) spores , and a diaphanopterodean nymph, probably Prochoroptera Handlirsch, with lycopsid spores in its gut (Kukalová-Peck 1987;Shear and Kukalová-Peck 1990). ...
... The coprolite record of palynivory is supplemented by rare occurrences of insect body fossils with gut contents from Mazon Creek in the Illinois Basin (Middle Pennsylvanian, Moscovian) Kukalová-Peck 1987;Shear and Kukalová-Peck 1990). These specimens provide direct evidence of lycopsid spore consumption across multiple insect lineages, including Eucaenus ovalis Scudder (Archaeorthoptera: Eucaenidae) with probable Lycospora spores (Carpenter and Richardson 1976;Scott and Taylor 1983), an unidentified archaeorthopteran with Cappasporites distortus Urban (Lepidodendrales) spores , and a diaphanopterodean nymph, probably Prochoroptera Handlirsch, with lycopsid spores in its gut (Kukalová-Peck 1987;Shear and Kukalová-Peck 1990). ...
... Coprolites from the Middle Pennsylvanian (early Moscovian) of Kentucky, USA, contain Lycospora spores , and coprolites from the late Moscovian of Iowa, USA contain megaspores of Sporangiostrobus Bode (Baxendale 1979). The gut of a probable Prochoroptera nymph (Diaphanopterodea) was filled with lycopsid spores (Kukalová-Peck 1987;Shear and Kukalová-Peck 1990), suggesting that some palaeodictyopteroid nymphs may have sought out food in the canopy. An arthropleurid with gut contents from the Middle Pennsylvanian contained lycopsid xylem (Rolfe and Ingham 1967;Rolfe 1969), and a massive coprolite from a Middle Pennsylvanian subhumid environment in the UK was filled with lycopsid particles, consistent with an arthropleurid producer (Falcon-Lang et al. 2015). ...
The Middle-Late Pennsylvanian Subperiod was marked by recurrent glacial-interglacial cycles superimposed on a longer-term trend of increasing aridity. Wetland and drought-tolerant floras responded with repeated migrations in the tropics, and a major plant turnover occurred in swamp ecosystems in parts of Euramerica near the Middle-Late Pennsylvanian boundary. However, the corresponding ecological and evolutionary responses of insects and other terrestrial arthropods are poorly understood. Here, we review the record of plant-arthropod interactions and analyse origination and extinction rates of insects during the Middle-Late Pennsylvanian. Although preliminary, plant-arthropod associations broadly persist through the Middle-Late Pennsylvanian boundary, and new damage types and host-plant associations first appear in the Late Pennsylvanian, possibly related to increased availability of accessible vascular and foliar tissues associated the shift from arborescent lycopsid to tree and seed fern dominance in Euramerican wetlands. Likewise, our analysis of the insect body fossil record does not suggest especially high rates of origination or extinction during this interval. Together, these results suggest that insects did not suffer major extinctions during the Middle-Late Pennsylvanian, despite short- and long-term changes in climate and environmental conditions.
Supplementary material at https://doi.org/10.6084/m9.figshare.c.6280586
... Although diplurans have an ancient origin that extends at least to the Devonian period (Grimaldi, 2010), their fossil record is extremely sparse given their often-diminutive size, delicate and lightly sclerotized bodies and a habitat preference that precludes fossilization. The oldest species recognizable as Diplura is Testajapyx thomasi Kukalová-Peck, 1987 from the Upper Carboniferous of Mazon Creek, IL, USA (Kukalová-Peck, 1987) although it differs from all Recent forms in a suite of morphological characters and has been discussed as a stem-group japygoid (Grimaldi, 2010). Other records consist of one species from the Lower Cretaceous Santana deposits of Brazil (Bechly, 2001;Wilson & Martill, 2001), three species from Upper Miocene or Pliocene onyx marble from Arizona (Pierce, 1950(Pierce, , 1951 and one species in Eocene Baltic amber (Silvestri, 1912). ...
... Although diplurans have an ancient origin that extends at least to the Devonian period (Grimaldi, 2010), their fossil record is extremely sparse given their often-diminutive size, delicate and lightly sclerotized bodies and a habitat preference that precludes fossilization. The oldest species recognizable as Diplura is Testajapyx thomasi Kukalová-Peck, 1987 from the Upper Carboniferous of Mazon Creek, IL, USA (Kukalová-Peck, 1987) although it differs from all Recent forms in a suite of morphological characters and has been discussed as a stem-group japygoid (Grimaldi, 2010). Other records consist of one species from the Lower Cretaceous Santana deposits of Brazil (Bechly, 2001;Wilson & Martill, 2001), three species from Upper Miocene or Pliocene onyx marble from Arizona (Pierce, 1950(Pierce, , 1951 and one species in Eocene Baltic amber (Silvestri, 1912). ...
Diplurans are among the earliest hexapods in the geological record. These primitively wingless relatives of insects are infrequently encountered despite being pervasive in soil habitats. Two groups have disparate adaptations for hunting, one mechanical and the other chemical, in Japygoidea and Projapygoidea, respectively. Here, we report three genera (two new) and four species of fossil Projapygidae preserved in Mesozoic and Cenozoic ambers: Electroprojapyx alchemicus gen. et sp. nov. in mid-Cretaceous Myanmar amber, and Symphylurinopsis punctatus gen. et sp. nov. and two species (unnamed) of Symphylurinus in Miocene Dominican amber. The exceptionally preserved specimens possess cerci that are morphologically specialized for expelling a glandular substance on prey. The new Cretaceous E. alchemicus is a stem group to all living species of the family and provides conclusive Mesozoic evidence for a rare type of predation involving offensive (vs. defensive) secretions. The specimen was fossilized in a preying-and-spraying position with its presumptive meal, a Symphypleona springtail (Collembola), reflecting the behavioural predatory repertoire of modern relatives. Apart from one Cenozoic species, these rare specimens represent the only fossil record of the family Projapygidae. Our findings demonstrate that these basal hexapods were predators of small arthropods of ancient forest soils and enrich our understanding of palaeoecological associations and behavioural strategies paramount for the survival of species.
... Other early groups are less clearly identifiable than Dasyleptus. A putative dipluran, Testajapyx thomasi (72), from the Pennsylvanian of Mazon Creek was tentatively considered to be related to stem-Dermaptera (165); some characters, such as the abdominal leglets, eversible vesicles, number of abdominal segments, and putatively entognathous mouthparts, were in doubt (66). Similarly, Ramsdelepidion schusteri from the Pennsylvanian of Mazon Creek was originally attributed to Zygentoma (72) but has been reconsidered as a stem group of Hexapoda (7,18,173). ...
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.
... A basal wingless insect from the suborder Monura (order Archaeognatha), illustrating proposed source tissues for wing origins. Modified and redrawn after(Kuklová-Peck, 1987). ...
Given the pervasiveness of gene sharing in evolution and the extent of homology across the tree of life, why is everything not homologous with everything else? The continuity and overlapping genetic contributions to diverse traits across lineages seem to imply that no discrete determination of homology is possible. Although some argue that the widespread overlap in parts and processes should be acknowledged as "partial" homology, this threatens a broad base of presumed comparative morphological knowledge accepted by most biologists. Following a long scientific tradition, we advocate a strategy of "theoretical articulation" that introduces further distinctions to existing concepts to produce increased contrastive resolution among the labels used to represent biological phenomena. We pursue this strategy by drawing on successful patterns of reasoning from serial homology at the level of gene sequences to generate an enriched characterization of serial homology as a hierarchical, phylogenetic concept. Specifically, we propose that the concept of serial homology should be applied primarily to repeated but developmentally individualized body parts, such as cell types, differentiated body segments, or epidermal appendages. For these characters, a phylogenetic history can be reconstructed, similar to families of paralogous genes, endowing the notion of serial homology with a hierarchical, phylogenetic interpretation. On this basis, we propose a five-fold theoretical classification that permits a more fine-grained mapping of diverse trait-types. This facilitates answering the question of why everything is not homologous with everything else, as well as how novelty is possible given that any new character possesses evolutionary precursors. We illustrate the fecundity of our account by reference to debates over insect wing serial homologues and vertebrate paired appendages. This article is protected by copyright. All rights reserved.
... Nonetheless, insect fossils before the Carboniferous are few; the Rhynie chert is followed by a window of 80 Ma (referred to as the 'hexapod gap') during which no insects are known [122]. The existence of pre-Devonian hexapods is a reasonable assumption, proposed already by early cladistic studies predating the molecular clock methodology [123]. Although a decade-old bounty of 1000 dollars has been put on an undisputable insect fossil from the pre-Devonian [124], this sum remains to be claimed. ...
Arthropods, the most diverse form of macroscopic life in the history of the Earth, originated in the sea. Since the early Cambrian, at least ~518 million years ago, these animals have dominated the oceans of the world. By the Silurian–Devonian, the fossil record attests to arthropods becoming the first animals to colonize land, However, a growing body of molecular dating and palaeontological evidence suggests that the three major terrestrial arthropod groups (myriapods, hexapods, and arachnids), as well as vascular plants, may have invaded land as early as the Cambrian–Ordovician. These dates precede the oldest fossil evidence of those groups and suggest an unrecorded continental “Cambrian explosion” a hundred million years prior to the formation of early complex terrestrial ecosystems in the Silurian–Devonian. We review the palaeontological, phylogenomic, and molecular clock evidence pertaining to the proposed Cambrian terrestrialization of the arthropods. We argue that despite the challenges posed by incomplete preservation and the scarcity of early Palaeozoic terrestrial deposits, the discrepancy between molecular clock estimates and the fossil record is narrower than is often claimed. We discuss strategies for closing the gap between molecular clock estimates and fossil data in the evolution of early ecosystems on land.
... The fact that three of the early branches of Pterygota have aquatic larvae has led to the idea that aquatic larvae may represent the ancestral state for the entire group (e.g., Kukalová -Peck 1978;1987;Shear and Kukalová -Peck 1990;Marden and Kramer 1994;Thomas et al., 2000;Zwick 2009) that became lost in further derived ingroups (at least twice). Yet, it has also been suggested that the aquatic larvae evolved independently in these lineages (e.g., Gullan and Cranston 2010;Bitsch 2012;Garwood et al., 2012) (Wipfler et al., 2019). ...
Aquatic larvae are known in three early branches of Pterygota: Ephemeroptera (mayflies), Plecoptera (stoneflies), and Odonata (dragonflies, damselflies). A common origin of these larvae has been suggested, yet also counterarguments have been put forward, for example, the different position of larval gills: laterally on the abdomen in Ephemeroptera, terminally in Odonata, variably in Plecoptera. We discuss recent fossil findings and report a new dragonfly-type larva from Kachin amber (Myanmar), which possesses ancestral characters such as a terminal filum, maintained in ephemeropterans, but lost in modern odonatan larvae. The new larva possesses lateral protrusions on the abdominal segments where in other lineages gills occur. Together with other fossils, such as a plecopteran retaining lateral gills on the abdomen, this indicates that lateral protrusions on the abdomen might have well been an ancestral feature, removing one important argument against the idea of an aquatic larva in the ground pattern of Pterygota.
Most paleontological textbooks deal with tracheates and chelicerates in only a cursory way because of their putatively poor fossil record. However, recent investigations into the paleobiology of these groups reveal that the fossil record is not only more extensive than previously assumed, but provides a wealth of information regarding both broad and detailed patterns of evolution of the two most diverse subphyla on the planet. Tracheata, including insects, entognaths and the various myriapod groups, are the most diverse subphylum. Insects alone are the most diverse class of animals known, outnumbering the combined species level diversity of all other animals. The Chelicerata, composed of the eurypterids, xiphosurids, arachnids and pycnogonids, are the second most diverse subphylum, with the diversity of arachnids exceeding all classes except for the insects. Consequently, not only does the evolution of tracheates and chelicerates provide an interesting story in itself, but these groups also provide us with insight into more general aspects of the evolutionary process that are of interest to the general evolutionary biologist as well as to the arthropod specialist.
Symmetries in the external world constrain the evolution of neuronal circuits that allow organisms to sense the environment and act within it. Many small “modular” circuits can be viewed as approximate discretizations of the relevant symmetries, relating their forms to the functions they perform. The recent development of a formal theory of dynamics and bifurcations of networks of coupled differential equations permits the analysis of some aspects of network behavior without invoking specific model equations or numerical simulations. We review basic features of this theory, compare it to equivariant dynamics, and examine the subtle effects of symmetry when combined with network structure. We illustrate the relation between form and function through examples drawn from neurobiology, including locomotion, peristalsis, visual perception, balance, hearing, location detection, decision-making, and the connectome of the nematode Caenorhabditis elegans.
Since the late Paleozoic, insects and arachnids have diversified in the terrestrial world so spectacularly that they have become unquestionably the most diverse group of organisms to ever inhabit the planet. In fact, this 300 million year interval may appropriately be referred to as the age of arthropods . What is the origin and history of terrestrial arthropods? How is arthropod diversity maintained on land? In this rhetorical context we will discuss (1) the degree to which terrestriality is found in arthropods, (2) the physiological barriers to terrestrialization that arthropod clades confronted, (3) the historical record of arthropod diversity on land based on paleobiological, comparative physiological and zoogeographical evidence, and (4) some tentative answers to the “why” of terrestrial arthropod success. We are providing a geochronologic scope to terrestriality that includes not only the early history of terrestrial arthropods, but also the subsequent expansion of arthropods into major terrestrial habitats.
Gigantic as well as very large mayflies from the middle Upper Carboniferous (Westphalian) strata of Europe and North America are described: the adult and nymph of Bojophlebia prokopi n. gen., n. sp. (Bojophlebiidae n. fam.) and the nymphs of Lithoneura piecko n. sp. and Lithoneura clayesi n. sp. (Syntonopteridae). Evolution of ephemerid wing venation during 300 million years is summarized. Autapomorphic, apomorphic, and plesiomorphic character states of venation are categorized. Venational nomenclature of Recent Ephemerida is emended based on its evolutionary changes. Evidence that wing veins occurred primitively as a pair of fluted sectors is documented in Carboniferous mayflies in the costa, subcosta, radius, anal, and jugal. Ephemeroids and odonatoids are sister groups that share the veinal anal brace AA fused with CuP at an area important for flight. Ancestral Odonatoephemerida are the sister group of the extinct haustellate Paleoptera. The Carboniferous nymphs bear three pairs of almost homonomous th...
It is generally agreed that Pennsylvanian age cyclothems in northern Illinois are deltaic in origin. However, there is little agreement on the specific application of the delta model. Appalachian and European cyclothems have been more successfully interpreted as they are generally thick, locally developed and often have close modern counterparts. A major difficulty in applying a delta model to the northern Illinois deposits lies in the widespread, continuous, yet thin development of the strata, particularly the coal members. To address this problem a cyclothem delta model is constructed and modified for static and fluctuating sea level conditions. Preliminary investigation of northern Illinois cyclic strata suggests a low gradient model (comparable to a tropical Mississippi Delta) coupled with generally static sea level conditions.
We examine several aerodynamic and thermoregulatory hypotheses about possible adaptive factors in the evolution of wings from small winglets in insects. Using physical models of Paleozoic insects in a wind tunnel, we explore the potential effects of wings for increasing gliding distance, increasing dispersal distance during parachuting, improving attitude control or stability, and elevating body temperatures during thermoregulation. The effects of body size and shape, wing length, number, and venation, and meteorological conditions are considered. Hypotheses consistent with both fixed and moveable wing articulations are examined. Short wings have no significant effects on any of the aerodynamic characteristics, relative to wingless models, while large wings do have significant effects. In contrast, short wings have large thermoregulatory effects relative to wingless models, but further increases in wing length do not significantly affect thermoregulatory performance. At any body size, there is a wing length below which there are significant thermoregulatory effects of increasing wing length, and above which there are significant aerodynamic effects of increasing wing length. The relative wing length at which this transition occurs decreases with increasing body size. These results suggest that there could be no effective selection for increasing wing length in wingless or short-winged insects in relation to increased aerodynamic capacity. Our results are consistent with the hypothesis that insect wings initially served a thermoregulatory function and were used for aerodynamic functions only at larger wing lengths and/or body sizes. Thus, we propose that thermoregulation was the primary adaptive factor in the early evolution of wings that preadapted them for the subsequent evolution of flight. Our results illustrate an evolutionary mechanism in which a purely isometric change in body size may produce a qualitative change in the function of a given structure. We propose a hypothesis in which the transition from thermoregulatory to aerodynamic function for wings involved only isometric changes in body size and argue that changes in body form were not a prerequisite for this major evolutionary change in function.
The origin and diversification of terrestrial communities has long been associated with the Devonian. The earliest known substantial assemblage of land plants and animals preserved in or close to their original habitats is still that of the Rhynie Chert peat bog, from Scotland, which is of Siegenian age.
Modern biotic provinces are centers of endemism surrounded by zones of coincidence of broad-ranged species. Provinces form coherent units of association above the community level. Superimposed ranges are used to construct a surface contoured for provincial diversity. Provincial boundaries overlap. Frequencies of provincial components along boundaries vary from season to season. Relict provinces survive in these zones. Biomeres, as chronologic provincial units, are transgressive and regressive, and exhibit complex intertonguing in boundary regions. Faunal and floral zonal sequences are replicable within the biomere, but are complicated by inversion and recurrence along boundaries. These anomalies are useful in locating boundary zones. Provinciality is due to homeostasis of the ecosystem and is proportional to diversity. Terrestrial biota is more provincial than marine biota. A biomere appears as a pioneer biota under new conditions. It differentiates about centers of habitat diversity, End_Page 615------------------------------ matures through increase in interdependence, and wanes to extinction under changing environment. Different styles of diversity are used to distinguish degree of provincial maturity. In Pennsylvanian and Lower Permian rocks, several terrestrial biomeres, based on insect-faunal and on floral ranges, are present across North America and Europe. One boundary is correlated with paleolatitude, another boundary is meridional. These biotically leaky boundaries are not associated with physiographic barriers. Zonation of terrestrial biomeres shows promise for fine-correlation of coal areas remote from standard and type stratigraphic sections. End_of_Article - Last_Page 616------------
The oldest paleontologically well-dated spore tetrads and cuti-clelike sheets of cells have been recovered from beds of Caradoc (mid-Ordovician) age from boreholes in the Murzuk (Murzuq) and Rhadames (Ghadamis) Basins of western Libya. These microfossils together with late Llandovery early Wenlock vascular plant megafossils (lycophytes and Psilophyton) known from Libya lead us to conclude that land plants, including vascular plants, probably had a long pre-Silurian record, extending at least into the basal Caradocian. Additonal evidence suggests that this record does not extend as far back as the earlier Late Cambrian (Dresbachian).