Sauropod teeth from the Upper Cretaceous Bissekty Formation of Uzbekistan

The isolated adult teeth of titanosaurian sauropods from the Upper Cretaceous Bissekty Formation at Dzharakuduk, Uzbekistan, differ little in overall structure but show considerable variation in enamel sculpturing and wear patterns. The crown shape of unworn juvenile teeth ranges from lanceolate to conical. Most specimens have enamel texture resembling crumpled paper or completely smooth enamel. Longitudinal grooves along the mesial and distal edges are present on only a few tooth crowns and might be developed on both the labial and lingual sides. Among 252 worn tooth crowns there are eight variants of wear patterns, all possible combinations of 0–2 apical and 0–2 lateral wear facets. The most common is wear pattern A1L0 (one apical facet, no lateral facets; 62.7%). The next most common variant has two apical and no lateral facets (A2L0, 12.3%). These apical wear facets include the primary wear facets, which are produced by an opposing functional tooth, and secondary wear facets, which are produced by a replacing upper tooth coming into contact with the functional lower tooth at a late wear stage. The relative abundance of tooth crowns with two apical wear facets possibly suggests incipient development of a tooth battery in the Bissekty titanosaur.     

Sauropod diversity in the Late Cretaceous of southwestern Europe: The lessons of odontology

Recent discoveries in southern France and northern Spain suggest that the morphology of titanosaurian teeth shows much greater variations that previously thought. It is suggested that the different morphotypes are informative at specific or generic level and that titanosaurian genera may indeed be recognized by their isolated teeth. It is also confirmed that juvenile titanosaurian teeth have a rather uniform, cylindrical morphology. Four different morphotypes are described for the Ibero-Armorican Island in the Late Cretaceous.     

Dentition and histology in titanosaurian dinosaur embryos from Upper Cretaceous of Patagonia,Argentina

Abstract: Exceptionally preserved sauropod embryos from the Late Cretaceous Anacleto Formation in Auca Mahuevo (Neuquén Province, Argentina) have provided fundamental information on titanosaurian ontogeny. This paper describes the dental composition, disposition and microstructure of the specimens. Embryonic teeth show size disparity, with lengths that vary from 1 to 3 mm and diameters ranging from 0.15 to 0.26 mm, with the most frequent length values between 2.5 and 3 mm. Apparently, a typical ‘pencil‐like’ tooth morphology and a dental formula of Pm 4, M 7–8/D10? remained constant during titanosaurian ontogeny, whereas the arrangement of teeth in the skull shows notable ontogenetic changes. Absence of wear facets on teeth suggests a lack of prenatal chewing movements. The enamel proportion is significantly higher in embryos than in mature titanosaurs, which suggests that this relationship varies during ontogeny. Embryonic bony tissue is composed of highly vascularized, cellular woven bone. The absence of osteonal tissue, the high degree of vascularization, the presence of numerous osteocytes and poor development of periosteal bone reveals that the Auca Mahuevo titanosaurs would have had a high early growth rate and that they were buried at a relatively advanced embryonic stage.     

Cheek tooth erosion in male babirusa (genus Babyrousa)

The wear on the occlusal surfaces of male babirusa cheek teeth was evaluated in 53 skulls of Babyrousa babyrussa from Buru and the Sula Islands and 87 skulls of B. celebensis from Sulawesi, Indonesia. Based on the comparative lengths of their continually growing maxillary canine teeth, the skulls were divided into five ‘age categories’ (A–E). Numerical and symbolic codes representing tooth wear were applied to each pillar (cusp region) of the mandibular and maxillary permanent third and fourth premolar teeth, and the first, second and third permanent molar teeth. There was no significant difference between the tooth wear patters of skulls in groups A and B, or in groups C and D, and so these were amalgamated. There was close correspondence in wear patterns between each side of the mouth in both species and in each age group. The wear patterns of the mandibular and maxillary teeth, although not identical, were very similar, as were the wear patterns of both species. In group A + B for both species tooth wear was relatively slight, with the M1 teeth experiencing most relative wear. There was almost no wear of the M3 teeth. In group C + D substantial wear of upper and lower M1 was evident. In group E more widespread wear of the cheek teeth was seen, with increased severity of M1 tooth wear, yet there was comparatively much less M2 and M3 tooth wear. The pattern of cheek tooth wear of the Babyrousa spp. was different from that shown by Sus scrofa. Differences in diet selection and processing were highlighted as potential contributing factors. The pattern of cheek tooth wear in male babirusa was not adequate for use to monitor their age.     

Specialized wear facets and late ontogeny in mammalian dentitions

Abstract

Many types of wear facets can be found in mammalian teeth. Some are related to the initial surface, others use the cross-section of the enamel as the main tool. In primary occlusal surfaces facets mark the gradual wear, that are related to a relatively late ontogeny. Facets in teeth with secondary occlusal surfaces, however, represent specific arrangements of crests of enamel and dentine. Such facets require some initial wear to become fully functional. The tooth morphology guarantees such facets to be effective for a long period of time. Therefore they can be discriminated as specialized facets. From the different types of facets three specialized ones were selected, blade facets, rasp-facets, and nipper-facets, because they ate widely distributed, function differently, and are comparable with mechanical tools. They are long lasting and differ in the amount of exposed dentine. The amount of dentine is used to differentiate phases during late ontogeny, the part of the life history, when teeth are exposed to wear. Consequently the various types of facets can be related to different ontogenetic phases. The relevant phases are prolonged at the cost of other ontogenetic phases. Therefore, the various specialized wear facets represent heterochronies within the ontogeny of teeth.     

Dietary adaptations of South African australopiths: inference from enamel prism attitude

The angle at which enamel prisms approach the wear surface holds information with regard to the stiffness of the tissue, as well as its wear resistance. Hence, analyses of prism orientation may shed light on questions of whether the thick enamel in hominins has evolved to confer stiffness or wear resistance to the teeth and may thus inform about the diet and behavioural ecology of these species. This was explored for Paranthropus robustus and Australopithecus africanus, whereby a distinction was made between prisms at the Phase I and Phase II facets. The results were compared with those obtained for Theropithecus, Macaca, and Potamochoerus for whom behavioural and/or experimental data are available, and were interpreted against simple mechanical principles. The South African hominins differ significantly in their relationships between wear facets and prism angulations. Teeth of P. robustus are better adapted to more vertical loads during mastication (Phase I), whereas those of A. africanus are better adapted to cope with more laterally-directed loads (Phase II) commonly associated with roll-crush and mastication. Overall, teeth of P. robustus appear stiffer, while those of A. africanus seem more wear resistant.     

Enamel hypoplasia in the deciduous teeth of great apes: variation in prevalence and timing of defects

The prevalence of enamel hypoplasia in the deciduous teeth of great apes has the potential to reveal episodes of physiological stress in early stages of ontogenetic development. However, little is known about enamel defects of deciduous teeth in great apes. Unresolved questions addressed in this study are: Do hypoplastic enamel defects occur with equal frequency in different groups of great apes? Are enamel hypoplasias more prevalent in the deciduous teeth of male or female apes? During what phase of dental development do enamel defects tend to form? And, what part of the dental crown is most commonly affected? To answer these questions, infant and juvenile skulls of two sympatric genera of great apes (Gorilla and Pan) were examined for dental enamel hypoplasias. Specimens from the Powell‐Cotton Museum (Quex Park, UK; n = 107) are reported here, and compared with prior findings based on my examination of juvenile apes at the Cleveland Museum of Natural History (Hamman‐Todd Collection; n = 100) and Smithsonian Institution (National Museum of Natural History; n = 36). All deciduous teeth were examined by the author with a ×10 hand lens, in oblique incandescent light. Defects were classified using Fédération Dentaire International (FDI)/Defects of Dental Enamel (DDE) standards; defect size and location on the tooth crown were measured and marked on dental outline charts. Enamel defects of ape deciduous teeth are most common on the labial surface of canine teeth. While deciduous incisor and molar teeth consistently exhibit similar defects with prevalences of ～10%, canines average between 70–75%. Position of enamel defects on the canine crown was analyzed by dividing it into three zones (apical, middle, and cervical) and calculating defect prevalence by zone. Among gorillas, enamel hypoplasia prevalence increases progressively from the apical zone (low) to the middle zone to the cervical zone (highest), in both maxillary and mandibular canine teeth. Results from all three study collections reveal that among the great apes, gorillas (87–92%) and orangutans (91%) have a significantly higher prevalence of canine enamel defects than chimpanzees (22–48%). Sex differences in canine enamel hypoplasia are small and not statistically significant in any great ape. Factors influencing intergroup variation in prevalence of enamel defects and their distribution on the canine crown, including physiological stress and interspecific dento‐gnathic morphological variation, are evaluated. Am J Phys Anthropol 116:199–208, 2001. © 2001 Wiley‐Liss, Inc.     

How far a protected area contributes to conserve habitat species composition and population structure of endangered African tree species (Benin,West Africa)

The Pendjari Biosphere Reserve located in the Sudanian zone of Bénin, is a protected area well managed, but mainly aimed at wild animal conservation. This study assessed its effectiveness to conserve habitat species composition and population structure of three endangered African tree species: Afzelia africana Sm., Pterocarpus erinaceus Poir. and Khaya senegalensis (Desv.) A. Juss. We randomly sampled 120 plots in the protected and surrounding unprotected habitats by inventorying plant species. For the three target species, we estimated adult and juvenile densities and recorded size classes. According to floristic composition four habitats groups were recognized in relation to human disturbance, vegetation type, and moisture. These were protected savannas, unprotected savannas, old fallows and gallery forests. The estimated adult densities of A. africana were similar between protected (14 ± 1.2 tree/ha) and unprotected savannas (17 ± 0.9 tree/ha) while for P. erinaceus the adult density was significantly higher in protected (12 ± 3.7 tree/ha) than in unprotected savannas (5 ± 1.9 tree/ha). Estimated adult density of K. senegalensis was also significantly higher in protected gallery forest (40 ± 5.8 tree/ha) than in unprotected one (29 ± 4.8 tree/ha). Juvenile densities of A. africana, K. senegalensis and P. erinaceus were higher in protected habitats than in unprotected ones but the difference was not significant. Skewness coefficient indicated that populations of investigated trees were declining in their protected habitats. However, in the case of A. africana and K. senegalensis populations seemed to be mostly threatened in the protected area. We concluded that although the studied protected area is effective to conserve some habitats species compositions, protection is not sufficient to guarantee future conservation of some threatened tree species.     

The dentition of a well-preserved specimen of <Emphasis Type="Italic">Camarasaurus</Emphasis> sp.: implications for function,tooth replacement,soft part reconstruction,and food intake

The basal macronarian genus Camarasaurus was the most common sauropod in the Upper Jurassic Morrison Formation of North America and is known from several complete and partial skeletons. The specimen used for this study is Camarasaurus sp. SMA 0002 from the Sauriermuseum Aathal, Switzerland. This specimen was found in the Howe-Stephens Quarry, Bighorn Basin, WY, USA. In this study, the dental morphology, characterized by the spatulate, broad-crowned teeth, the tooth replacement pattern, and the function of the dentition and its implications for food intake is described. Features such as the absence of denticles, the wrinkled pattern of the enamel, and the occurrence of large wear facets on older teeth are characteristic for Camarasaurus sp. A slab of sediment with soft tissue impressions ranging up to the middle part of the crown suggests the presence of a gingival soft tissue structure partially covering the teeth. The wrinkled enamel on the crown of the teeth of Camarasaurus sp. and other sauropods is interpreted as indication of this cover of gingival connective tissue. In addition, there possibly was a keratinous beak, which together with the gingiva held the teeth in the jaw and provided stability for teeth in which the root is almost completely resorbed.     

Response to Delibes-Mateos et al. : Pellet size matters

In Rueda et al. [Rueda, M., Rebollo, S., Gálvez-Bravo, L., 2008. Age and season determine European rabbit habitat use in Mediterranean ecosystems. Acta Oecol. 34, 266–273] we used a threshold of 6 mm faecal pellet diameter to differentiate between adult and juvenile European rabbit (Oryctolagus cuniculus) habitat use. Delibes-Mateos et al. designed a housing experiment with 12 adult rabbits and criticised the choice of 6 mm as a threshold to separate adult and juvenile rabbit pellets, claiming that adults can produce pellets both larger and smaller than 6 mm in similar proportions. In response to their criticism we argue the following. The selection of a 6 mm threshold has a bibliographic basis, it is not a new method developed by Rueda et al. and produces consistent results when applied in the field. Assuming that Delibes-Mateos et al. results are accurate, we should have found a greater number of <6 mm pellets than >6 mm, overall and seasonally, which is not the case. We believe that the use of commercial pelleted food, keeping animals isolated in small cages for over a year, and the use of adult rabbits only, makes the experimental design used by these authors not suitable to refute the usefulness of separating rabbit pellets smaller and larger than 6 mm diameter as indicators of changes in the relative abundance of juvenile and adult rabbits in the field. Finally, we agree with the authors that the use of indirect methods of animal aging would require case-specific validation studies; however, we believe these studies should be correctly designed.     

Antimicrobial activity of medicinal plants and induction of defense related compounds in banana fruits cv. Robusta against crown rot pathogens

A total of 72 plant extracts were tested in vitro for their ability to inhibit the mycelial growth of Lasiodiplodia theobromae and Colletotrichum musae the causal agents of crown rot disease of banana. The results showed that the leaf extract of Zimmu (an interspecific hybrid of Allium cepa L. × Allium sativum L.) and tuber extract of Zehneria scabra recorded maximum inhibition of mycelial growth and spore germination of both the test pathogens. The dipping of banana fruits in Zimmu leaf extract at 25% conc. exhibited 100% inhibition of crown rot disease in cold storage (14 °C) up to 35 days and increased the shelf life to 64 days. However, at room storage (28 ± 2 °C), the same treatment exhibited 86% inhibition of crown rot disease up to 12 days. It was found that the treatment of banana fruits with Zimmu leaf extract did not alter the organoleptic properties of banana. The biochemical analysis of banana fruits treated with Zimmu leaf extract showed significant increase in phenylalanine ammonia-lyase (PAL), chitinase and β-1,3-glucanase activities and enhanced accumulation of phenolic compounds compared to other treatments. These findings suggest that the effect of Zimmu leaf extract on crown rot disease may be associated with the direct fungi toxic property against the test pathogens and elicitation of defense related compounds in banana fruits.     

New material of Diacodexis (Mammalia,Artiodactyla) from the early Eocene of Southern Europe

Diacodexeidae are the first representatives of Artiodactyla in the fossil record. Their first occurrence is at the very base of the Ypresian (earliest Eocene, 56.0 Ma) with Diacodexis, a genus well diversified during the early Eocene in Europe, especially during the MP7–MP8 + 9 interval. However, most of European species are documented by scarce material, retrieved from single localities. In this work, we describe new Diacodexis material from ～MP7 and ～MP8 + 9 localities of Southern Europe, including material of D. antunesi from Silveirinha, considered as the most primitive European Diacodexis species, and material from three localities from Southern France (Fordones, Palette, and La Borie). The new material documents Diacodexis premolar morphology and deciduous dentition which bear potentially important phylogenetic information, as well as astragali, including a specimen from Silveirinha that constitutes the earliest occurrence of an astragalus of the genus Diacodexis in the European fossil record. Investigation of the enamel microstructure reveals that early European species had a simple enamel pattern with one-layered Schmelzmuster composed of ‘basic’ radial enamel only, instead of the two-layered Schmelzmuster (thin radial enamel + thick layer of Hunter-Schreger bands) observed on North American species and so far considered to represent the primitive condition within Artiodactyla. In accordance with previous studies, our observations highlight that Diacodexis gigasei from Belgium is morphologically closer to the North American species D. ilicis than to D. antunesi from Portugal. The latter species, together with D. aff. antunesi from Fordones, appears to be morphologically closer to the Asiatic taxa D. indicus and D. pakistanensis. Finally, we found numerous similarities between D. cf. gigasei from Palette and D. gigasei, a result that challenges the intra-European provincialism that characterizes the earliest Ypresian. Diacodexis gigasei could be one of the rare species shared by the northwestern and southwestern European bioprovinces.     

Late Quaternary woolly mammoth (Mammuthus primigenius Blum) remains from southern Transdanubia,Hungary

Six samples of subfossil tusk, bone and tooth remains from the woolly mammoth (Mammuthus primigenius Blum) were discovered in south-western Hungary. The remains are relatively well preserved in a Late Pleistocene loess deposit. The samples have been radiocarbon dated (AMS) and are of Late Weichselian (MIS 2) age (21.8–24.1 ka cal BP). The skull fragments, the tusks and maxillary teeth are in close proximity to associated postcranial remains, indicating that the mammoth died where it was found. The size and characteristics of skeletal elements have allowed us to determine that this was a mature male of about 38 years of age.     

Three new freshwater species of centrohelid heliozoans: Acanthocystis crescenta sp. nov., A. kirilli sp. nov., and Choanocystis minima sp. nov.

Three new species of centrohelid heliozoans Acanthocystis crescenta, A. kirilli, and Choanocystis minima from two freshwater lakes of Valamo Island (North-Western Russia) were studied using light- and scanning electron microscopy. The main apomorphy of A. kirilli are slightly branched radial scales. Acanthocystis crescenta has characteristic radial scales with crescent-like structures on their tips. Cell diameter of these two species does not exceed 10 μm. Radial scales of C. minima have truncated tips and lack apical teeth; plate scales of this species are oviform, without lateral notches or central rib. The latter organism is the smallest heliozoan species known; the diameter of its body is about 3 μm. The possible domination of small heliozoans in the cryptic diversity of centrohelids is discussed.     

A human parietal fragment from the late Early Pleistocene Gran Dolina-TD6 cave site,Sierra de Atapuerca,Spain

The TD6-2 level of the Gran Dolina cave site (Sierra de Atapuerca, Burgos, Spain) has yielded an assemblage of about 170 human fossil remains dated to > 800 ka (probably MIS 21) and assigned to the species Homo antecessor. In this study, we describe for the first time a large portion of a parietal bone (ATD6-100/168). The morphology of the fractures on the bones is compatible with a peri-mortem trauma. The superior parietal areas are flat. There is a large parietal foramen and one smaller accessory parietal foramen. Middle meningeal vessels are not particularly developed, but they are distributed in both anterior and posterior districts, with the parietal vasculature originating from the posterior branch. The meningeal vessels show multiple minor connections with the pericranial and diploic vascular systems. The diploe is not particularly developed, and large diploic channels are not detected. The bone is thin when compared with adult fossil humans, and equivalent to juvenile values. All these characters suggest that the parietal ATD6-100/168 probably belonged to a juvenile individual, with plesiomorphic endocranial traits similar to those described for H. ergaster/erectus. The derived temporal, maxillary and dental traits in Homo antecessor and the primitive parietal morphology further point to distinct (mosaic) patterns of morphological evolution of face and braincase.     

SEM study of the oral cavity of members of the Kyphosidae and Girellidae (Pisces,Teleostei), with remarks on Crenidens (Sparidae), focusing on teeth and taste bud numbers and distribution

The present study compares dental morphology and taste bud distribution in the oral cavity of four species of the teleost family Kyphosidae, five species of Girellidae and one species of Sparidae. Some of these species are predominantly herbivorous, while others are omnivorous with a high portion of invertebrates in their diets. All the kyphosids feature one row of frontal cusped teeth on their jaws and areas of miniature teeth on the tongue and palate; the girellid species feature 2–5 rows of denticulate teeth, and no teeth inside the oral cavity. The total number of taste buds in the oral cavity is higher in Girella spp. than in the kyphosids, and their number is species-specific, not correlated with fish size. For example, Kyphosus bigibbus of 500 mm standard length bears 1780 taste buds, while the kyphosid Neoscorpis lithophilus of 80 mm bears 3460. The maximum number of taste buds, 7900, is found in Girella punctata of 60 mm standard length, and the minimum number, 1320, in Kyphosus vaigiensis of 175 mm. The higher number of taste buds appears to characterize species that possess a more diversified diet. The present study contributes to our understanding of the differences between the two families Kyphosidae and Girellidae. In particular, it relates the eco-morphological adaptations to the type of diet consumed by the various species.     

Hunter-Schreger band orientation in interproximal enamel of modern human molars with complex morphological traits

A recent paper concerning characteristic groove-shaped wear in interproximal attritional facets of Neandertal posterior teeth correlated groove formation with deep enamel structural traits (Villa & Giacobini, 1995). Further studies were carried out on modern human teeth in order to evaluate a possible correlation between deep enamel structure and some crown complex morphological features. Seventy-five isolated molars from medieval and recent ossuaries as well as five molars recently extracted for therapeutic purpose were examined using scanning electron and optic microscope ground sections of interproximal enamel. The results showed possible correlations between deep enamel structural characteristics (i.e., Hunter-Schreger band verticalization) and some crown complex traits (i.e., occlusal groove pattern, mesial and distalfovea, mesial and distal accessory tubercules, interproximal marginal ridges). Observations reported in this study showed a significantly higher frequency of Hunter-Schreger band verticalization in lower molars with an occlusal complex pattern (i.e. primitive fissure pattern “Y” or “+”). This observation suggests a positive correlation among crown complex traits (which occur frequently in Neandertal teeth) with deep enamel structural characteristics. In addition, analysis of teeth grouped according to individual showed crown morphological traits and deep enamel structural characteristics corresponding to the same individual. This observation supports a genetic basis in occlusal shape and microstructural dental traits.     

Pennsylvanian fusulinids and calcareous algae from Sonora (northwestern Mexico), and their biostratigraphic and palaeobiogeographic implications

Pennsylvanian carbonates are widespread in Sonora (Mexico) and contain a diverse biota of foraminifers and calcareous algae. Detailed studies here are devoted to the outcrops of the Sierra Agua Verde and Cerro El Tule. The Late Atokan (early Late Moscovian part), Desmoinesian (= late Late Moscovian) and Missourian (= Kasimovian) stages are especially rich in fusulinids and algae. The principal zones of fusulinids of Wilde encountered are A3, DS1 and MC1–2. New data are given about the genera Fusulinella, Parawedekindellina, Zellerella, Komia and Paraepimastopora, in order to establish migrations or vicariances between Mexico and Palaeotethys.     

Palaeoenvironmental significance of Miocene larger benthic foraminifera from the Xisha Islands,South China Sea

We studied 344 samples from Well XK-1 in Xisha Islands, South China Sea, and identified 66 species of larger benthic foraminifera, providing critical evidence for biostratigraphy and palaeoenvironmental interpretation of the Miocene reef carbonate sequence. Three assemblages are recognized, namely, Spiroclypeus higginsi–Borelis pygmaeus Assemblage (Letter Stage Te5, Early Miocene, 1256.28–1180.15 m), Nephrolepidina–Miogypsina Assemblage (Tf, Middle Miocene, 1031.10–577.04 m), and Cycloclypeus–Heterostegina Assemblage (Tg, Late Miocene, 468.13–380.42 m). On the basis of the palaeoecological preference of the larger foraminifera, we interpret that the Miocene carbonate sequence was deposited mainly in a warm tropical shallow water environment, characterized by five stages of continuous long-term evolution: backreef lagoon to shelf in the Early Miocene, normal to frontal reef in the early Middle Miocene, backreef lagoon to shelf in the later Middle Miocene, normal to frontal reef in the early Late Miocene, and proximal forereef shelf in the later Late Miocene.     

Evaluation of Pseudomonas syringae strain ESC-11 for biocontrol of crown rot and anthracnose of banana

Pseudomonas syringae strain ESC-11 and 250 μg/ml each of thiabendazole (TBZ) and imazalil reduced crown rot of banana caused by Fusarium aff. sacchari by 30–36% and 83–86%, respectively, in laboratory experiments. Four field trials performed in Costa Rica varied in treatment combinations. In field trials 1 and 2, 125 and 250 μg/ml each of TBZ and imazalil + 0.5% or 1% alum (aluminum ammonium sulfate) and ESC-11, and 250 μg/ml each of TBZ and imazalil + 1% alum reduced rot and mold. ESC-11 alone or with 0.5% alum significantly reduced rot and mold in field trial 2. In trial 3, 50 and 100 μg/ml of TBZ alone and with ESC-11 reduced mold. In trial 4, 125 μg/ml each of TBZ and imazalil and ESC-11, and 300 μg/ml each of TBZ and imazalil reduced rot, and 50 and 125 μg/ml each of TBZ and imazalil and ESC-11, and 300 μg/ml each of TBZ and imazalil reduced mold. In three field trials, there was no significant difference among treatments for latex staining. In field trial 2 only, combinations of TBZ, imazalil, and alum with or without ESC-11, reduced anthracnose, caused by Colletotrichum musae. The complex of crown rot fungi, order of treatment application, effect of alum and fungicides on ESC-11, concentration of ESC-11, and level of disease may contribute to the variation in crown rot and anthracnose control by ESC-11. Though ESC-11 alone was not effective in reducing disease, further testing in combination with low rates of fungicide should be done.