RETRODISPLACEMENT OF THE ORAL AND ANAL OPENINGS IN DENDRASTERID SAND DOLLARS

In regular echinoids, the mouth opening (or peristome) and the anal opening (or periproct) are located centrally. In irregular echinoids, the peristome tends to shift toward the anterior end of the test, whereas the periproct typically shifts toward the posterior end. This produces an anatomically polarized morphology, which is consistent with functional expectations. In the dendrasterid sand dollars, however, the peristome and periproct have been displaced in the “wrong” directions; the peristome has shifted posteriorly, whereas the periproct has shifted anteriorly. These movements, which run counter to functional expectations, may be termed “retrodisplacements.” This study presents a new model for the development of the oral surface in dendrasterids. The model assumes that the “Dendraster pattern” of development (which occurs in dendrasterids) was derived from the older “Echinarachnius pattern” (which occurs in other northern Pacific sand dollars). The Echinarachnius pattern includes two successive phases of asymmetric growth: the first phase favors anterior growth, whereas the second phase favors posterior growth. These two phases are normally balanced, and at maturity, the test appears to be symmetrical. But if the second phase of unequal growth were suppressed through a heterochronic change in development, the effects on test development would be profound. On the aboral surface, the predicted effects include posterior displacement of the apical system. On the oral surface, the predicted effects include retrodisplacement of the peristome and periproct, as well as conspicuous changes in ambulacral and interambulacral development. In fact, all of the predicted effects are characteristic features of dendrasterids. The model assumes that the oral and aboral surfaces could be simultaneously affected by the same developmental processes. This assumption is supported by empirical evidence: in dendrasterids, there is a strong correlation between the displacement of the apical system (on the aboral surface) and the displacement of the peristome (on the oral surface). The displacement of the apical system (known as “apical eccentricity”) is regarded as a valuable adaptation, because it facilitates suspension feeding. But if oral and aboral development are linked, then selective pressure for apical eccentricity would simultaneously produce new oral characters as well. Thus, the retrodisplacement of the peristome and periproct in dendrasterids may be related to developmental constraints. These unusual characters may have little or no functional significance.     

Loriolella,a key taxon for understanding the early evolution of irregular echinoids

Loriolella Fucini is an enigmatic and poorly understood Early Jurassic sea-urchin that has been interpreted both as a regular and an irregular echinoid. The discovery of new and well-preserved material of the type species L. ludovicii (Meneghini) from the Upper Pliensbachian (Lower Jurassic) of Lombardy, Italy, has clarified the morphology of this genus for the first time. Loriolella resembles regular echinoids in having a large (almost certainly monocyclic) apical disc enclosing the periproct, and large primary interambulacral tubercles and spines. However, its peristome is extremely small with only vestigial buccal notches and auricles, and the ambulacra expand adorally forming cassiduloid-like phyllodes. Cladistic analysis suggests that Loriolella is the earliest known microstomatid irregular echinoid. Its unique mixture of primitive and derived characters is important for interpreting the initial steps by which irregular echinoids arose from diadematoid-like regular ancestors.     

Morphological similarity between echinoid taxa as a result of convergent and parallel evolution

Convergence is observed in groups, which are phylogenetically remote. A flat test is typical for some representatives of the order Cassiduloida, such as Jurassic and Early Cretaceous species of the genera Pygurus and Clypeus, and also many Cenozoic «sand dollars,» i.e., echinoids of the order Clypeasteroida. Both usually inhabit coarse sandy grounds of shallow areas. The superorder Spatangacea includes the so-called Echinocorys life form, which is characterized by an oval test with superficial nonpetaloid or subpetaloid ambulacra, marginal or inframarginal periproct, and absence of fascioles. These are the following genera: Early Cretaceous Corthya (family Collyritidae), Late CretaceousLate Paleocene Echinocorys (family Holas-teridae), Paleocene Isaster, Recent Isopatagus (family Isasteridae), Recent Scrippsechinus (family Palaeotro-pidae), and Recent Urechinus (family Urechinidae). In contrast to the majority of spatangaceans with the burrowing mode of life, these genera dwell on the substrate surface. In the Cenozoic, the monobasal apical system appears in some genera of the order Spatangoida, most genera of the order Cassiduloida, all groups of the orders Clypeasteroida and Oligopygoida, and in the genus Echinoneus (order Holectypoida). The paral-lelism is revealed in groups connected by remote relationships. At the end of the Middle Jurassic (Callovian) and, especially, Late Jurassic, the so-called disasterid echinoids (superorder Spatangacea) show a distinct trend to the loss of contact between ocular plates I and V and apices of the posterior ambulacra with the periproct, which are shifted to the anterior part of the apical system (genera Collyrites, Collyropsis, Cyclolam-pas). At the same time, the peristome of some genera was displaced to the anterior margin of the test, which became bilaterally symmetrical in outline. However, in the Jurassic, all spatangaceans remained disasterid echinoids, i.e., had a disjunct apical system, which can be interpreted as a somewhat “abnormal” state. This trend disappeared only at the beginning of the Cretaceous, when “normal” forms with a joint apical system appeared, that is, the families Holasteridae (genera Eoholaster and Holaster, order Holasteroida) and Toxas-teridae (genus Toxaster, order Spatangoida). Interesting examples of synchronous parallelism are provided by the appearance of meridosternous (diasternal) plastron in two collyritid genera (Tetraromania and Corthya) in the Barremian, whereas in the holasterid genus Holaster, this type of plastron apparently appeared in the Valanginian (heterochronous parallelism). The ethmolitic type of the apical system appeared at the end of the Cretaceous and Paleocene at least in five families: Schizasteridae, Paleopneustidae, Brissidae, Spatangidae, and Loveniidae.     

Ecophenotypic Variation and Developmental Instability in the Late Cretaceous Echinoid Micraster brevis (Irregularia; Spatangoida)

The Late Cretaceous echinoid genus Micraster (irregular echinoids, Spatangoida) is one of the most famous examples of a continuous evolutionary lineage in invertebrate palaeontology. The influence of the environment on the phenotype, however, was not tested so far. This study analyses differences in phenotypical variations within three populations of Micraster (Gibbaster) brevis from the early Coniacian, two from the Münsterland Cretaceous Basin (Germany) and one from the North Cantabrian Basin (Spain). The environments of the Spanish and the German sites differed by their sedimentary characteristics, which are generally a crucial factor for morphological adaptations in echinoids. Most of the major phenotypical variations (position of the ambitus, periproct and development of the subanal fasciole) among the populations can be linked to differences in their host sediments. These phenotypic variations are presumed to be an expression of phenotpic plasticiy, which has not been considered in Micraster in previous studies. Two populations (Erwitte area, Germany; Liencres area, Spain) were tested for stochastic variation (fluctuating asymmetry) due to developmental instability, which was present in all studied traits. However, differences in the amount of fluctuating asymmetry between both populations were recognised only in one trait (amount of pore pairs in the anterior paired petals). The results strengthen previous assumptions on ecophenotypic variations in Micraster.     

Heterochrony and heterotopy in the phylogeny of sea urchins

The role of heterochrony is evident in ontogeny and phylogeny of irregular (exocyclic) sea urchins. After metamorphosis, a juvenile passes the stage of regular (endocyclic) sea urchin, in which the periproct is surrounded by plates of the apical system. A shift of the periproct in the area of the fifth interambulacrum occurs in extant taxa at early stages of postlarval development and is accompanied by the reduction of genital plate 5. In some ancient (Jurassic) adult irregular sea urchins, the endocyclic state of the apical system is retained for a long time and the derivative of the fifth genital plate is sometimes observed even in Early Cretaceous species. Considerable transformations in the structure of the lower test surface in members of the order Spatangoida are manifested in changes in the relative positions of plastron plates and ambulacral areas I and V, separation of sternal plates from the labrum, etc. The mechanism of these changes is connected with translocation or “sliding” of sutures of particular plates as a result of nonuniform growth and partial resorption. The study of evolutionary lineages of Cretaceous and Cenozoic sea urchins has shown that the evolution was connected with the directional changes in some morphological characters at late ontogenetic stages. The process was accompanied by either extension, peramorphosis (lineages of the genera Micraster, Infulaster–Hagenowia in the European Province), or the loss of these stages, paedomorphosis (Hemiaster (Bolbaster) lineage, Late Eocene–Middle Miocene of Australia). The phenomena of heterochrony and heterotopy in the development of peripetal, marginal, and lateroanal fascioles in the Late Cretaceous and Paleocene families Hemiasteridae, Schizasteridae, and Paleopneustidae are described. The heterotopy is also illustrated by the example of the development of additional genital pores on ocular plates II and IV of the Middle Jurassic species Pygomalus analis (Disasteroida); its apical system has five pores instead of four. In the Late Cretaceous species Guettaria roccardi (Holasteroida), ocular plates II and IV have two pores each; in the apical system, there are eight genital pores instead of four. In some members of the order Holectypoida, the place of lost genital plate 5 is occupied by a new plate sometimes pierced by a pore, but judging from crystallographic data, it is not homologous to other genital plates. The order Clypeasteroida is characterized by the development of very small pores in both ambulacral and interambulacral fields; they provide passage for numerous accessory tube feet.     

Shallow Traces (Pits) in the Test of the Irregular Echinoid Echinocorys scutata Leske from the Chalk (Upper Cretaceous) of the United Kingdom

Specimens of epifaunal irregular echinoids in the Upper Cretaceous of northern Europe have been reported with patterns of circular, nonpenetrative parabolic pits, Oichnus paraboloides (Bromley), in the apical region. Specimens of Echinocorys scutata Leske from the Chalk at two sites in southeast England were commonly penetrated by this trace, generated by an indeterminate pit-forming organism. Pits commonly surround the apical system and, less commonly, occur within it; they occur preferentially anteriorly. Pits occur within plates, not along margins or sutures. Crosscutting of pits indicates that multiple spatfalls probably occurred. The host echinoid added new test plates adjacent to the apical system; thus, plates bearing O. paraboloides were moved abapically. The reduction in number of pits away from the apex, including those with echinoid tubercles reestablished on the base, indicates that, following death of an infester, the echinoid “reclaimed” and infilled them with calcite. The pit-former was most probably an unmineralized invertebrate that used E. scutata as a domicile which provided good access to food-rich currents for suspension feeding. Although the systematic position of the pit-former is unknown, similar infestations are known from other Upper Cretaceous echinoids and Mississippian crinoids.     

IRREGULAR ECHINOIDS—AN INSUFFICIENTLY KNOWN GROUP

Some aspects of echinoid evolution from an endocyclic towards an exocyclic apical system, in the light of embryological and palaeontological data, are discussed. A primitive feature—imbrication of interamhulacral plates in Jurassic irregular echinoids, recently described—proves once more that this group is insufficiently known. Every revision of taxonomic or phylogenetic concepts concerning this group of echinoids should be preceded by careful morphological analysis of the material.     

Variabilité architecturale du test chez le genre Collyrites (Echinoidea, Disasteroidea) au Jurassique moyen

The aim of this paper is to study the evolution of the architecture of the « apical system–periproct complex » of the genus Collyrites (Echinoidea, Disasteroidea) for species from the Bathonian and Callovian stages (Middle Jurassic). The studied material comes from several localities of the Paris basin. The apical system of the genus Collyrites is subdivided in two parts: (i) an anterior part composed by four genital (1, 2, 3, 4) and three ocular (II, III, IV) plates, called trivium; (ii) a posterior part composed by two ocular plates (I, V) and one genital plate (5), without gonopor, which circle the periproct, called bivium. The genital plate 5, which may collapse in the periproct, is not always visible. The Bathonian–Callovian transition is marked by a subdivision of the bivium in two parts: the periproct breaks up from the posterior ocular plates (I, V). These morphological changes are associated with architectural modifications. The trivium stays relatively stable during the Bathonian and the Callovian, but a supplementary plate may be inserted into the trivium. The bivium shows important modifications linked to the separation of the periproct and the ocular plates (I and V). Such a separation is marked by a strong development of supplementary plates, these ones keeping in connection with the periproct and the ocular plates I and V. The supplementary plates are more and more developed whereas the distance between the periproct and ocular plates increases. The connection between the trivium and the bivium is similarly provided by supplementary plates. The size of these plates seems to significantly increase between the Bathonian and the Callovian. Moreover, some specimens from the Bathonian and the Callovian may have an atypical architecture with a supplementary genital plate or a genital plate with two pores. The “extraxial axial theory” allows to recognize two types of skeleton: (i) an “axial skeleton” corresponding to ocular plates and plates of the ambulacra and interambulacra; (ii) an “extraxial skeleton” corresponding to the genital and supplementary plates, and the periproct. Architectural modifications between the Bathonian and the Callovian is a result of a more important development of the “extraxial skeleton” while the “axial skeleton” shows few modifications during this time interval.     

Early post-metamorphic ontogenesis of deep-sea spatangoids (Echinoidea, Spatangoida) of the NE Atlantic Ocean

Abstract. The early juvenile development of 3 species of irregular deep-sea echinoids of the NE Atlantic was examined using scanning electron microscopy. Although the post-metamorphic development of these 3 spatangoids is very similar, the morphology and development of fascioles (specialized bands of heavily ciliated minute spines) facilitates distinguishing the species. In Hemiaster expergitus and Spatangus raschi , the fascioles present in the early juvenile develop to form adult fascioles. In Brissopsis lyrifera , a juvenile fasciole of unknown function disappears during ontogenesis, and new adult subanal and periproct fascioles arise. Fascioles of S. raschi and B. lyrifera are present at a very early stage of development (∼0.7 mm test length), whereas in H. expergitus the fasciole appears much later (∼3 mm test length). Animals smaller than 0.8 mm test diameter have not yet developed a functional mouth and are still considered as endotrophic postlarval stages. Mouth development is similar for all spatangoids examined.     

Development of the nervous system in the brittle star Amphipholis kochii

There are several studies of neural development in various echinoderms, but few on ophiuroids, which develop indirectly via the production of pluteus larvae, as do echinoids. To determine the extent of similarity of neuroanatomy and neural development in the ophiuroids with other echinoderm larvae, we investigated the development of the nervous system in the brittle star Amphipholis kochii (Echinodermata: Ophiuroidea) by immunohistochemistry. Immunoreactive cells first appeared bilaterally in the animal pole at the late gastrula stage, and there was little migration of the neural precursors during A. kochii ontogeny, as is also the case in echinoids and holothuroids. On the other hand, neural specification in the presumptive ciliary band near the base of the arms does occur in ophiuroid larvae and is a feature they share with echinoids and ophiuroids. The ophiopluteus larval nervous system is similar to that of auricularia larvae on the whole, including the lack of a fine network of neurites in the epidermis and the presence of neural connections across the oral epidermis. Ophioplutei possess a pair of bilateral apical organs that differ from those of echinoid echinoplutei in terms of relative position. They also possess coiled cilia, which may possess a sensory function, but in the same location as the serotonergic apical ganglia. These coiled cilia are thought to be a derived structure in pluteus-like larvae. Our results suggest that the neural specification in the animal plate in ophiuroids, holothuroids, and echinoids is a plesiomorphic feature of the Ambulacraria, whereas neural specification at the base of the larval arms may be a more derived state restricted to pluteus-like larvae.     

Physical forces experienced by echinoid eggs in the oviduct during spawning: comparison of the geminate pair Echinometra vanbrunti and Echinometra lucunter

During spawning, eggs of echinoids are released from the gonad and traverse a narrow oviduct-gonopore complex before being released into the water surrounding the spawning female. As the eggs traverse the oviduct-gonopore complex, a velocity gradient develops within the fluid formed by the eggs. This velocity gradient imposes a shear stress on the eggs that ranges from a maximal value at the walls of the oviduct-gonopore complex and declines linearly to zero at the center of the complex. In addition to shear stress, the eggs of some echinoids also experience strain. Strain is imposed on the eggs in situations where the diameter of the egg is greater than the diameter of the oviduct-gonopore complex. Both shear stress and strain have the potential to damage echinoid eggs during spawning and are likely selective pressures acting on the physical properties of protective extracellular layers, which surround the eggs of these organisms. The amounts of shear stress and strain experienced by eggs of a geminate pair of echinoids, Echinometra vanbrunti and E. lucunter, are calculated. Calculations of shear stress based on the mean spawning rates and dimensions of the oviduct-gonopore complexes of specimens collected during this study ranged up to 30 Pa for E. vanbrunti and up to 17 Pa for E. lucunter. To obtain conservative estimates of shear stress, these calculations were repeated incorporating a spawning rate that was reduced by one order of magnitude below the mean rate measured for both species. These calculations yielded shear stresses that ranged up to 13 and 6 Pa for E. vanbrunti and E. lucunter, respectively. The diameter of the oviduct-gonopore complex of echinoids increases linearly with the diameter of the tests of the adults. Because the diameter of the oviduct-gonopore complex influences the amount of shear stress experienced by eggs of echinoids, shear stress calculations were repeated incorporating the full range of potential diameters of the oviduct-gonopore complexes from data in the literature. Using the mean spawning rates obtained from each species, the amount of shear stress experienced by eggs ranged up to 32 Pa for E. vanbrunti and up to 398 Pa for E. lucunter. When the mean spawning rates were reduced by one order of magnitude, the amount of shear stress experienced by E. vanbrunti eggs ranged up to 14 Pa and up to 140 Pa for the eggs of E. lucunter. The eggs of E. vanbrunti do not experience strain during spawning because the diameter of the its eggs is smaller than the diameter of its oviduct-gonopore complex at all stages of adult growth. In contrast, the diameter of the eggs of E. lucunter is larger than the diameter of the oviduct-gonopore complex at small adult sizes and the eggs of this species experience true strains ranging up to 1.2. These results indicate that there are substantial differences in the amounts of physical forces that the eggs of echinoids experience during spawning both within species at different stages of adult growth and between species. Different amounts of shear stress and strain experienced by eggs among echinoid species may select for differences in the mechanical properties of protective extracellular layers that surround them.     

SYMMETRY,LOCOMOTION, AND THE EVOLUTION OF AN ANTERIOR END: A LESSON FROM SEA URCHINS

Bilaterally symmetrical, “regular” sea urchins in the Family Echinometridae (Class Echinoidea; Phylum Echinodermata) were found to lack a locomotor anterior. Heterocentrotus mammillatus and Echinometra mathaei were observed while locomoting. Members of both ellipsoidal species were found to proceed with their short or long axis foremost with statistically equivalent frequencies. This finding demonstrates that the evolution of bilateral symmetry is not always accompanied by the evolution of a locomotor “anterior” end. The elliptical echinometrid sea urchins provide a particularly appropriate study group for investigating the relationship between the evolution of body form and locomotor behavior. Although the radially symmetrical regular sea urchins, from which the echinometrids sprang, lack a locomotor anterior, all “irregular” echinoids, which are also derived from a regular ancestor but are bilaterally symmetrical, possess an “obligate” locomotor anterior. The symmetry and behavior exhibited by the elliptical echinometrid sea urchins therefore demonstrates that the first irregular echinoids (which exhibit bilateral symmetry by definition) need not have possessed a locomotor anterior as they do today.     

Revision of <Emphasis Type="Italic">Scoliocystis</Emphasis> (Rhombifera: Echinoencrinitidae) and related cystoid genera

Species attributed to Scoliocystis Jaekel, 1899, including the type species S. pumila (Eichwald, 1860) and S. thersites Jaekel, 1899 from the Ordovician of the Leningrad Region, are reviewed. Scoliocystis sp. from the Upper Ordovician of Estonia, figured by Hecker (1964), is redescribed as Maennilocystis heckeri gen. et sp. nov. and it is attributed to the family Callocystitidae Bernard, 1895. The genus Scoliocystis is intermediate between the families Cheirocrinidae and Echinoencrinitidae in having five periproct border plates (as in cheirocrinids), but lacking plate R5 and having a reduced oral area, as in echinoencrinitids, but is retained in the Echinoencrinitidae. The similarity of Scoliocystis to the unusual North American cheirocrinid genus Sprinkleocystis is convergent. The genus Gonocrinites Eichwald, 1840, with two species, is restored among echinoencrinitids, as having four periproct border plates, whereas Echinoencrinites sensu stricto has three. The families Echinoencrinitidae and Callocystitidae arose from cheirocrinids with closed plate circlets by the loss of plate R5 and substitution of radial:lateral for radial:radial pectinirhombs. In addition, echinoencrinitids are characterized by a reduced oral area, whereas in callocystitids, the recumbent ambulacra are extensive. Scoliocystis is a stem-group genus to Echinoencrinitidae plus Callocystitidae. Several glyptocystitoid genera independently developed advanced pectinirhombs.     

Host selection,host-use pattern and competition in Dissodactylus crinitichelis and Clypeasterophilus stebbingi (Brachyura: Pinnotheridae)

The genera Dissodactylus and Clypeasterophilus are commensal or parasitic crabs that live in association with irregular sea urchins. Only a single pinnotherid species is expected to occur on a sand dollar; however, during field observations, D. crinitichelis and Clypeasterophilus stebbingi were found co-occurring on a single host (Clypeaster subdepressus). Laboratory experiments and in situ observations were conducted to explore mechanisms of host selection and use by C. stebbingi, as well as the possibility of competition with D. crinitichelis. Analysis of 213 specimens of Encope emarginata and 33 C. subdepressus revealed a high frequency of infestation (90 %) by brachyurans. The number of D. crinitichelis was positively correlated with the size of E. emarginata. During field observations and experiments, D. crinitichelis was able to colonize both echinoids, while C. stebbingi was found mainly on C. subdepressus. No exclusion was reported between individuals of the same sex for both crab species. However, D. crinitichelis may be territorial, since a single male occurred when adult females were added to the host. D. crinitichelis and C. stebbingi co-occurred on the same individual of C. subdepressus on rare occasions during field observations. C. stebbingi may exclude D. crinitichelis from the echinoid to avoid competition, as observed during the experiments. We suggest a species-specific association between C. stebbingi and C. subdepressus, while D. crinitichelis associated mainly with E. emarginata, but also with C. subdepressus. Both crabs may harm the echinoids by consuming their spines, but additional research is necessary to determine whether this is a parasitic association.     

Un modèle d'ontogenèse: La croissancede Toxaster granosus kiliani (Lambert)), échinide spatangoide du Valanginien

A complete study of the ontogeny of the irregular urchin Toxaster granosus kiliani (Lambert)) was achieved from an important lower cretaceous sample. In a first time, ontogenic variations of each character were analysed versus length chosen as size reference. This first step had shown that very important changes in shape take place during the development of the young urchin: increase in height, downward migration of the periproct, forward movement of the peristome… In a second time, a growth crisis was proved and described. Such a crisis occurs, from 12 mm to 16 mm length, as a momentary acceleration of the growth (i.e. the rate of growth increases during a short time). The reality of this crisis was demonstrated by both mathematical and biological ways and also by comparing with other actual and fossil populations. The whole of these results leads to a synthetical pattern of growth divided into three main stages which are successively a juvenile, a preadult and an adult stage. This division was proved also by a principal components analysis.     

A novel DNA polymerase homologous to Escherichia coli DNA polymerase I from a higher plant, rice (Oryza sativa L.)

A novel DNA polymerase, designated as OsPolI-like, has been identified from the higher plant, rice (Oryza sativa L. cv. Nipponbare). The OsPolI-like cDNA was 3765 bp in length, and the open reading frame encoded a predicted product of 977 amino acid residues with a molecular weight of 100 kDa. The OsPolI-like gene has been mapped to chromosome 8 and contains 12 exons and 11 introns. The encoded protein showed a high degree of sequence and structural homology to Escherichia coli pol I protein, but differed from DNA polymerase γ and θ. The DNA polymerase domain of OsPolI-like showed DNA polymerase activity. Subcellular fractionation analysis suggested that the protein is localized in the plastid. Northern and western blotting, and in situ hybridization analyses demonstrated preferential expression of OsPolI-like in meristematic tissues such as shoot apical meristem, root apical meristem, leaf primordia and the marginal meristem. Interestingly, no expression was detected in mature leaves, although they have a high chloroplast content. These properties indicated that OsPolI-like is a novel plant DNA polymerase. The function of OsPolI-like is discussed in relation to plastid maturation.     

Two new species of Panorpa (Mecoptera,Panorpidae) from the Hengduan Mountains in Yunnan,China

Two new species of the genus Panorpa Linnaeus, 1758 are described from the Hengduan Mountains in Yunnan, China. Panorpa diqingensis sp. n. can be distinguished from its congeners by vertex dark brown, and occiput yellow; pleura dark brown; gonocoxites bearing a cluster of stout black setae on inner apex; parameres S-shaped in male genitalia. Panorpa dispergens sp. n. can be readily differentiated from other Panorpa species by apical band with a hyaline spot in wings; gonocoxites with a cluster of stout dark brown setae and 3–5 long setae on inner apex; female medigynium with a pair of basal plates. The number of Panorpa species in Yunnan is raised to six. A key to Yunnan species of Panorpa is presented.