2 Окт 2012 Gurisar 5
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Relevant to the current discussion is the holotype of Aneimites austrina Etheridge Jr. However, in our opinion, the specimens under consideration and those described by Morris exhibit sufficiently distinct and diagnostic features that they should be excluded from B. Instead, these specimens should be considered as a new taxon, apparently confined to Australia. To avoid an indeterminate designation for these specimens in the present article, we are adopting here the binomen of Morris , namely Triphyllopteris austrina Etheridge Jr.
Herbivorized and unherbivorized T. For additional comparison, unherbivorized pinnae of T. Co-occurring with T. The fern-like foliage has been interpreted as representing pteridosperm seed fern plants Morris , , ; Rigby , , , and they are the dominant elements; by contrast, sphenopsid and lycopsid material occur as subordinate elements Morris ; R.
Based on recent examination of these specimens by R. These plant taxa also support an early Serpukhovian date for the deposit. Three specimens of T. These specimens represent fragments of ultimate-order pinnae that bear small, alternately arranged pinnules, also illustrated in unherbivorized specimens Fig. The rachises of T. The pinnules are ovate to obovate, sessile, decurrently inserted on the rachis, and they display lateral margins that are entire while the distal margins are broadly rounded, semilobate and diminutively crenulate.
Pinnule size extends to 16 mm in length by 8. The venation is dense, open, and dichotomizes several times before reaching the distal margin. The laminae are apparently thick. External feeding of P. These digital photos are a continuation of documentation using traditional photographic methods provided in Fig.
Another excised edge is at the center of d, in a deep sinus between adjacent pinnules, illustrated in g, with wispy veinal stringers, one of which is indicated by an arrow. A third example of folivory d , at lower left, is enlarged in h, characterized by a relatively thin reaction rim and a thin flap of necrotic tissue projecting into the matrix. The last example in d, at upper-right, is an extensive excision with major subordinate cuspules, enlarged in i, and exhibiting a veinal stringer arrow that parallels overall venational trend.
Comparison of the details of extant orthopteran margin feeding with the pattern of folivory on T. The overall trajectory of feeding progresses from right to left straight arrow , although each individual feeding event is represented by a cuspule forming a localized, semicircular path in the opposite direction, from left to right curved arrow.
Additionally, note veinal stringers along the eaten margin signifying unconsumed vascular tissue. A frontal view of a grasshopper c , exhibiting general relationships between hypognathous, mandibulate mouthparts and a consumed grass leaf, an example of modern margin feeding Gangwere For these four examples of P. Two examples of P. Enlargements e and f illustrate the inferred path of overall cuspate feeding straight arrow and localized feeding trajectories of several cuspule feeding events curved arrows.
Fossil plant material of Late Mississippian age was examined by R. This material was originally recovered by various collectors during the late s to early s. Specimens were examined macroscopically under a Wild stereomicroscope, and the photographs in Fig. Photographs in Fig. Separate overlay drawings are provided in Figs. Length and width measurements were taken of the major cuspate excisions and the much smaller cuspules forming the scalloped edges, which are listed in Table 1.
Herbivory was demonstrated by 1 a characteristic C- or U-shaped, stereotyped style of excision with subordinate cuspules; 2 distinctive reaction tissue probably callus along the damaged margin; and 3 the projection of veinal stringers and truncated vascular tissue defining the boundaries between adjacent cuspules within the larger cuspate excision, and as flaps of necrotic tissue Labandeira ; Fig.
These features are distinctly different from abiotic types of foliar damage Wilson , Vincent , Katterman Dimensions of P. Herbivore damage of this specimen Figs. This damage type is equivalent to DT12 in the damage-type system of Labandeira et al. The documented stratigraphic range of this type of feeding formerly was Middle Pennsylvanian van Amerom and Boersma to Pliocene Straus , but the range now has been extended earlier to the earliest Late Mississippian this study; Fig.
Herbivorized specimens: AMF Figs. Present are six, moderately preserved, discontinuous, oval-elongate excisions of the foliar margin, ranging from 0. Within the larger excisions are mostly an uninterrupted series of smaller, subordinate cuspules ranging in size from 0. These cuspules are often separated by projecting veinal stringers or truncated vascular tissue Fig. Distinct reaction rims of variably thickened scar tissue line the six excisions Fig. The large-scale removal of foliar tissue is interpreted herein as a result of extensive cuspate margin-feeding activity Coulson and Witter Clear reaction-tissue rims are evidenced by the thickened and darkened edge surrounding the damaged foliar margin, which may be attributable to callus tissue.
The presence a scalloped edge along the incised foliar margin of most of the cuspate excisions suggests repeated backward rotations of the head of a margin-feeding orthopteroid or related and similarly feeding insect Fig. Each rotation represents a feeding event that would result in a cuspule on a foliar margin and eventually be equivalent to bolus sections in an insect fecal pellet or coprolite found in Pennsylvanian-age deposits Labandeira and Phillips This type of feeding is similar to that described for extant orthopteroid insects by Gangwere and Kazikova , which range from plant host generalists to specialists Isely , Otte and Joern , Jones and Firn , Rowell et al.
A modern orthopteroid mode of feeding is illustrated in Fig. These six C- or U-shaped hemispherical excisions occur along the margin of a richly veined pteridosperm pinnule margin and are characterized by a thickened and often scalloped rim of reaction tissue, thus providing strong evidence that this feature has resulted from the consumption of live foliar tissue.
This plant damage is consistent with a generalized, folivorous orthopteroid insect with strong, mandibulate mouthparts both in terms of overall shape of P. However, the temporal placement of this trace fossil is 6 m. Thus, P. A temporal disjunction occurs between the earliest documented appearance of distinctive plant damage with that of a delayed body-fossil record of potential culprits.
Such a temporal disjunction is a recurring theme throughout the fossil record—one that contrasts a temporally older plant—insect associational record with a relevant but younger body-fossil record that can supply culprit taxa Scott et al. A much less likely culprit is a diplopod myriapod Zherikhin Rare body fossils of diplopods and other litter inhabiting and ground dwelling myriapod taxa occur from the Devonian to the Mississippian—Pennsylvanian boundary Wilson and Shear , Wilson et al.
Given this context, it is important that Mississippian triphyllopteroid-type fronds such as T. Based on evidence from the inferred elevated stature of the T. Although folivory is now documented for the Late Mississippian earliest Serpukhovian , and the earliest documented leaves megaphylls are rare occurrences from the Early Devonian Fig. This type of external herbivory indicates that mandibulate microarthropods had the capability for chewing or otherwise abrading live, three-dimensional plant organs such as erect stems before or during the initial appearance of leaves.
This damage contains reaction tissue, including callus, which indicates consumption of live stem tissue. A culprit for such damage remains unknown, although mandibles from an earlier arthropod, Rhyniognatha hirstii Hirst and Maulik , have been redescribed by Engel and Grimaldi , and based on its mouthparts, may be consistent with exophytic plant consumption also see Labandeira These mandibles originate from the earlier Pragian Rhynie Chert of Scotland, and the deposit contains evidence for the consumption of inner parenchymatic tissues and sporivory as indicated by coprolites Habgood et al.
There is a quantum increase in the abundance of laminate leaves in floras worldwide, commencing during the Early Visean, approximately at — Ma Fig. However, herbivore associations throughout the Mississippian are poorly known. Most documented arthropod-mediated damage involves detritivory Chaloner et al. One interesting occurrence is a zygopterid fern, Symplocopteris wyattii , from the older mid Tournasian of Queensland, Australia, which bears roots and aphlebiae that contain clusters of microcoprolites within chambers of evacuated tissue, probably bearing reaction tissue indicating herbivory Hueber and Galtier These aphlebiae are linear, appendiculate structures on stems that, unlike leaves, are supplied by one vascular strand, and they have highly three-dimensional but not planated cross sections.
The fossil record of folivory, until now, extended to the early Middle Pennsylvanian Moscovian of equatorial Euramerica van Amerom ; van Amerom and Boersma ; Scott and Taylor ; Scott et al. A notable absence of evidence for folivory characterizes the Early Pennsylvanian Bashkirian to early Moscovian. For the Middle Pennsylvanian Moscovian, there are many occurrences. Evidence of herbivory from this interval consists principally of three-dimensional plant damage from British coal-ball material Seward ; Jeffrey ; Stopes ; Holden , , ; De Witt West , and it is contrasted to considerably fewer examples from the compression or impression record e.
Folivory becomes more widespread, albeit still at relatively low levels, during the late Moscovian, particularly in the Illinois Basin, and historically has been best demonstrated for the medullosan seed fern compression taxon Macroneuropteris scheuchzeri Hoffmann Cleal et al. From taphonomically very different, carbonate permineralized coal-ball deposits, foliar herbivory apparently picks up during the Late Pennsylvanian Kasimovian in peat substrated, coal-swamp forests of the Illinois and other Euramerican basins.
By the middle Artinskinan of the Early Permian, the type and level of folivory increases dramatically in Euramerica, but in extrapaludal, mostly fluvially associated habitats characterized by mineralic substrates and mesic vegetation such as those described floristically by DiMichele et al. This damage includes skeletonization, surface abrasion, and intercostal hole-feeding where large swaths of lamina are removed between secondary veins.
This type of herbivory also is reproduced on true gigantopterid taxa in younger late Early Permian Kungurian and early Middle Permian Roadian deposits in Cathaysia Glasspool et al. By contrast, another, older Sakmarian site from Texas records a less diverse spectrum of herbivore damage types and levels of herbivory Labandeira and Allen The Late Permian crisis, linked to the reduction of pteridosperm-dominated flora worldwide, had a major effect on all types of herbivorous plant—insect associations, reflected in the virtual absence of insect-mediated damage in the few known Early Triassic floras Labandeira , ,.
The earliest documented occurrence of external foliage feeding, now recorded for the Late Mississippian, provides for a discussion of three associated and salient issues. They are 1 the issue of major ecological lags between the origin of land plant leaves and their subsequent consumption by herbivores; 2 the contrast between older plant—insect associational data versus younger body-fossil data that may identify culprits for the older associations; and 3 the emerging, distinctive pattern of pteridosperms as the favored plant host for Paleozoic folivores.
Leaves in vascular plants megaphylls independently originated four times Boyce and Knoll , although additional origins are possible, depending on various resolutions of early land-plant phylogeny Boyce a. The earliest leaf-like structures were simple linear structures microphylls borne by the Lycopsida during the latest Silurian immediately before Ma, which became common in Early Devonian floras Bonamo et al.
The sister-group to the Lycopsida, the Euphyllophytopsida, diversified during the subsequent Devonian, producing four major clades, each of which convergently produced megaphyllous leaves that were laminate and multiveined. The four lineages within the Euphyllophytopsida that convergently evolved structurally similar leaves beginning in the Early Devonian and expanding during the Middle and Late Devonian were, from oldest to youngest, 1 progymnosperms, 2 sphenopsids, 3 ferns, and 4 seed plants Boyce a,b ,.
Earlier representatives of these clades bore leafless, photosynthetic stems with stomata and subepidermal airspaces analogous to foliar mesophyll tissue Edwards The earliest known leaf is Eophyllophyton bellum Hao and Beck Hao and Beck , a progymnosperm of the mid-Early Devonian Pragian of Yunnan, China; by contrast, the earliest herbivorized leaf is the seed plant pteridosperm T. This represents a lag of 76 m.
However, current evidence, including this report, indicates that folivory was a delayed strategy. Three reasons for the overall lack of folivory among numerous Mississippian and Early Pennsylvanian leaf flora were mentioned by Scott and colleagues First, folivory appeared late because it was an advanced and difficult dietary habit for terrestrial arthropods and thus its chronological delay is expected also see Southwood Second, most leaves appeared relatively late in terms of major vascular plant structural innovations, and they became diverse during the later Carboniferous, although it is now evident that the origin of leaves occurred iteratively throughout the Devonian and Mississippian Boyce and Knoll And third, specimens with folivory have yet to be found, and thus their delay is more apparent than real.
Given this context, especially the first point above, Southwood proposed three different reasons from an understanding of insect physiology that account for the Paleozoic delay in herbivory. Given the preeminent role of secondary compounds determining current foliage palatability Price , Schowalter , some have proposed that secondary compounds initially were a by-product of metabolic biosynthetic pathways that were either passively produced or alternatively used for physical protection, such as flavonoids for warding off UV light absorption Cooper-Driver Subsequently, as herbivores became ecologically important, secondary compounds became actively selected as a plant antiherbivore defenses to reduce food palatability and nutrient assimilation rates, such as the crucial role that phenolics provide in herbivore deterrence Feeny , Bernays et al.
There is evidence that some Paleozoic seed plant lineages, such as medullosans, were producing noxious secondary compounds Bergen et al. With regard to the cooptation of secondary compound defenses, Beerling et al. Thus, planated megaphyll leaves became more abundant among Late Devonian and Mississippian flora, despite their remote origin during the Early Devonian Hao et al.
This potential food resource nevertheless may have been sufficiently rare during this interval that folivory was not a preferred feeding strategy, explaining the prolonged temporal lag. A related issue is first-occurrence data involving earlier trace-fossils and relevant later body-fossils in the assessment of plant-insect associations and their responsible culprits.
Although this issue has been discussed previously Scott et al. A germane example from the plant-insect associational record is the temporal disjunction between the presence of leafminer damage on Early Cretaceous angiosperm leaves and the significantly younger and relevant lepidopteran body-fossil record Kozlov , Labandeira et al. For the present case of folivory and their likely insect herbivores, a temporal disjunction of 6 m.
The alternative hypothesis that a diplopod was responsible for this damage would require major shifts in understanding of the feeding roles and life habits of Paleozoic diplopods, an unlikely prospect. Primitively flightless fossil hexapods occur in Early and Middle Devonian deposits.
However, these Devonian fossils are collembolans and archaeognathans Hirst and Maulik ; Shear et al. It is not until the Mississippian to Pennsylvanian boundary Ma; Brauckmann et al. Put another way, because there are no known hexapods from the interval ranging from the Upper Devonian to the Mississippian—Pennsylvanian boundary, there is a 6-m.
If the occurrence of the earliest Late Mississippian P. A more taxonomically circumscribed, orthopteroid culprit clade could have been true Orthoptera, which have their earliest occurrence in the late Middle Pennsylvanian Sharov and would represent an even greater time gap of 17 m. An important aspect about the occurrence of folivory on Australian T.
Together with ferns, seed plant foliage represents a greater complexity in venational pattern than for previous leaf originations Boyce b. Triphyllopteroid foliage is such an example, and has been referred to the pteridosperms by many authors Skog and Gensel , Meyen , Taylor and Taylor , Vega and Archangelsky Although Mississippian age fronds of Triphyllopteris -type from the Laurasian Realm have been included in the family Calamopityaceae Taylor , Meyen , Taylor and Taylor , sterile fronds referred to Triphyllopteris from the mid-Carboniferous latest Mississippian to Early Pennsylvanian Gondwanan Realm can be related to another austral family of lyginopterid affinity, the Austrocalyxaceae Vega and Archangelsky Notably, coeval Mississippian pteridosperm reproductive structures seeds and ovulate organs found throughout Gondwana resemble most closely the Lyginopteridaceae Morris , Erwin et al.
Thus, given the apparent absence of calamopityacean remains in the Carboniferous deposits of Gondwana and the fact that triphyllopteroid fronds were probably common to more than one plant family, T. Many of these clades are distantly related Rothwell and Serbet , Hilton and Bateman , although they collectively are referred to a group with an excess of plesiomorphic over apomorphic characters when compared with Mesozoic and Recent seed plants.
In addition to structures such as medullosacean resin rodlets interpreted as chemical defenses Bergen et al. By comparison, minimal folivory has been documented for co-occurring cordaitalean, coniferalean, cycadophyte and various pteridophyte clades Labandeira Collectively, this pattern for the origin and subsequent spread of folivory indicates that of the four vascular-plant clades that evolved leaves during the Devonian and earlier Mississippian—progymnosperms, sphenopsids, ferns, and seed plants—it was the seed plants as pteridosperms that became overwhelmingly dominant hosts for folivores.
This pattern of the expansion of folivory occurred globally, although the timing differs in Laurasia mostly Euramerica , Gondwana, and Cathaysia. In Euramerica, folivory is recorded minimally in equatorial wetlands during the Middle Pennsylvanian, increased significantly in the Late Pennsylvanian Labandeira , and was followed during the Permian by expansion principally onto different pteridosperm clades in the fluvial environments of dryer regions with mineralic soils Beck and Labandeira , Labandeira and Allen By contrast, in Gondwana folivory commenced during the later Mississippian, but it was disrupted subsequently by either glaciation or the secondary effects of periglacial environments during the Pennsylvanian and earliest Permian Gastaldo et al.
This was followed by renewed folivory throughout the ensuing Permian across the supercontinent Plumstead ; Srivastava ; McLoughlin a,b , ; Adami-Rodrigues and Iannuzzi Little is known of Paleozoic folivory for Cathaysia, although limited occurrences of folivory are documented for the mid-Permian Glasspool et al. The trend of preferential use of pteridosperm hosts by folivores persisted to the end of the Paleozoic in Laurasia, Cathaysia and Gondwana Labandeira and Allen , and it has continued on more derived lineages of seed plants to the present day, albeit in modified form in spite of more recent colonization of pteridophytes by varied insect herbivores Balick et al.
Five conclusions are drawn from this study of early insect folivory and an analysis of the earliest records of other types of folivory. These conclusions have implications regarding the early evolution of folivory among some of the oldest, well-documented terrestrial ecosystems. The oldest documented occurrence of insect-mediated folivory in terrestrial ecosystems consists of cuspate excisions of P.
However, a 6-m. The origin and subsequent expansion of Paleozoic folivory is repeatedly associated with seed plants, overwhelmingly pteridosperms, rather than other seed plant clades and three other clades of vascular plants—progymnosperms, sphenopsids, and ferns—from which leaves have independently originated. Trace fossil data from plant-insect associations can provide valuable evidence for documenting insect feeding patterns that are unavailable from the relevant body fossil record.
We are very grateful to Robert Jones for access to R. Thanks are extended to Finnegan Marsh, who rendered the figures; Marcie Harder took digital photos for Fig. We are grateful to three reviewers who commented on earlier versions of this manuscript. Adami-Rodrigues , K and R Iannuzzi. Acta Geol. Leopold 24 : Google Scholar. Permian plant-insect interactions from a Gondwana flora of southern Brazil. Fossils Strata 51 : a. Paleont 7 : b. Lethaia 29 : Insect-fern interactions: macrolepidopteran utilization and species-area association.
Entomol 5 : Biochemical and evolutionary aspects of arthropod predation on ferns. Oecologia Berl. Palaeobotanist 28—29 : Bot 80 : Beck , C B Current status of the Progymnospermopsida. Palynol 21 : Early Permian insect folivory on a gigantopterid-dominated riparian flora from north-central Texas.
Palaeoecol : Nature Lond. Paleoenvironmental contexts and taphonomic modes. Terrestrial ecosystems through time. University of Chicago Press. Chicago, IL. Unusual resin chemistry from Upper Carboniferous pteridosperm resin rodlets. American Chemical Society symposium based on amber, resinite and fossil resin.
American Chemical Society. Washington, DC. Herbivores and plant tannins. Res 19 : Leclercqia, Haskinsia , and the role of leaves in delineation of Devonian lycopod genera. Gaz : Boyce , C K The evolutionary history of roots and leaves. Vascular transport in plants. Amsterdam, The Netherlands. Boyce , C K Patterns of segregation and convergence in the evolution of fern and seed plant leaf morphologies.
Paleobiology 31 : b. Evolution of developmental potential and the multiple independent origins of leaves in Paleozoic vascular plants. Paleobiology 28 : The stratigraphical position of the oldest known Pterygota Insecta, Carboniferous, Namurian. Belg : Paleontol 12 : Ameghiniana 34 : Fossil evidence for plant-arthropod interactions in the Palaeozoic and Mesozoic. B : Chandra , S and K J Singh. Plant fossils from the type locality of Talchir Formation and evidence of earliest plant-animal activity in Gondwana of India.
Ninth International Gondwana Symposium. Hyderabad, India, Oxford, United Kingdom. A revised taxonomy for Palaeozoic neuropteroid foliage. Taxon 39 : Cooper-Driver , G A Biological roles for phenolic compounds in the evolution of early land plants. Gensel , P G and D Edwards , editors. Plants invade the land: evolutionary and environmental perspectives. Columbia University Press. New York. Forest entomology: ecology and management. Wiley Interscience. Crawley , M J Herbivory: the dynamics of animal-plant interactions.
University of California Press. Berkeley and Los Angeles, CA. Buenos Aires. The Carboniferous Period. A geologic time scale— Cambridge University Press. Cambridge, United Kingdom. Root ecology. Hist 5 : Palaios 15 : Paleontol 78 : On Paleozoic plants from marine strata: Trivenia arkansana Lyginopteridaceae gen. Bot 90 : Edwards , D Cells and tissues in the vegetative sporophytes of early land plants. New Phytol : Palaeontographica B : New light shed on the oldest insect. Early seed plants in the Southern Hemisphere: I.
Associated ovulate and microsporangiate organs from the Carboniferous of Peru. Palynol 80 : Feeny , P Plant apparency and chemical defense. Biochemical interaction between plants and insects. Gangwere , S K Relationships between the mandibles, feeding behavior, and damage inflicted on plants by the feeding of certain acridids Orthoptera. Entomol 1 : Out of the icehouse into the greenhouse: a late Paleozoic analog for modern vegetational change.
Today 6 : Genise , J F Ichnotaxonomy and ichnostratigraphy of chambered trace fossils in palaeosols attributed to coleopterans, ants and termites, pp. IcIlroy [ed. Publ : Foliar herbivory in Late Palaeozoic Cathaysian gigantopterids. Palynol : Geologic time scale —why, how, and where next. Lethaia 37 : Plant-insect interactions and coevolution during the Triassic in western Europe. Evidence for an early terrestrial food web: coprolites from the Early Devonian Rhynie Chert.
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The Geochron displays the following: Legal Zone Time, Greenwich mean time, Greenwich apparent time, local apparent time, local mean time, moment of sunrise, moment of sunset, duration of daylight, sun's meridian passage, sun's equation of time, degrees latitude, degrees longitude, the geographic extent of the prevailing day and date.
The basis of daily time measurement is the relationship of the earth's surface to the sun. Thus, when the earth makes one revolution about its axis relative to the sun, it is said to be one day. However, the speed of the earth's travel around the sun varies so that over a period of one year, some days are shorter and others longer, causing the relative zenith, or noon position of the sun to vary accordingly.
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One's location in longitude within a time zone also affects the relationship between "apparent noon" and "legal zone time noon". For example, if the international dateline coincides with the "mean sun" mean noon , half of the earth's surface is today and the other half is tomorrow or yesterday, depending on which half one's position is.
At this point of time, the same day exists around the earth. In practice, however, legal zone boundaries are highly irregular and non-conforming. Further distortion of "ideal" zone time occurs where daylight savings time is used. It also varies due to the changing speed of the earth along its orbit around the sun.
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