Morphology and ontogeny of the eodiscoid trilobite Sinodiscus changyangensis from the lower Cambrian of South China

Abstract: A large number of complete specimens together with numerous disarticulated sclerites of the eodiscinid trilobite Sinodiscus changyangensis Zhang in Zhou et al., 1977 have been collected from the lower Cambrian Shuijingtuo Formation in Changyang, Hubei Province, South China. An ontogenetic series is established based on the immature and mature exoskeletons including the previously unknown protaspides and meraspides, in particular. No further substages can be differentiated in the protaspid specimens herein. Changes that took place during the meraspid period include the addition of postcephalic segments and prominent pygidial larval notches in early meraspid development which became progressively less distinct and disappeared in degree 2. Two holaspid stages are recognized based on the addition of a new pygidial segment, indicating that the start of the holaspid phase preceded the onset of the epimorphic phase and accordingly, its developmental mode is attributed to the protarthrous pattern. The trunk segmentation schedule of S. changyangensis is discussed, which is similar to other primitive eodiscoid trilobites, that is, as the boundary between the thorax and pygidium migrated posteriorly, there is no change in the number of the trunk segments. The processes of liberation of the thoracic segment and segment insertion into the pygidium are separated from one another, and the two different mechanisms, somitogenesis and tagmosis, progress independently during the ontogenetic development of the postcephalic region of these primitive eodiscinids.     

Intraspecific variation of trunk segmentation in the oryctocephalid trilobite Duyunaspis duyunensis from the Cambrian (Stage 4, Series 2) of South China

A detailed exploration of growth and trunk segmentation of the oryctocephalid trilobite, Duyunaspis duyunensis Chang & Chien in Zhou et al. 1977, from the lower Cambrian (Stage 4, Series 2) Balang Formation in western Hunan Province, South China, is presented. Because of the excellent preservation, the complete post‐protaspid ontogenetic series from merapsid degree 0 to the holaspid phase is described. The ontogenetic series reveals new information on morphological changes such as the migration of the posterior branch of the facial sutures (from proparian to opisthoparian) and contraction of the posteromedial notch in the pygidium. The abundance of articulated specimens available from a narrow stratigraphical interval makes this material singularly useful for studying the morphogenesis and post‐embryonic growth of D. duyunensis in comparison with other oryctocephalids. Strong evidence that multiple numbers of pygidial segment are recognized in each meraspid degree as well as in the holaspid period showed unusual intraspecific variability in the rate of trunk segmentation, providing insights into how Cambrian subisopygous trilobites controlled their body patterning, including size, shape and trunk segment number in both thorax and caudal plate during growth.     

Articulation,interlocking devices and enrolment in Monkaspis daulis (Walcott, 1905) from the Guzhangian,middle Cambrian of North China

Monkaspis daulis (Walcott 1905 ) is the first enrolled trilobite to be documented from the Kushan Formation (Guzhangian, Cambrian Series 3) of Northern China. It has a wide pygidium that allowed a sphaeroidal enrolment type to be achieved, covering the cephalon and anterior trunk segments. Monkaspis daulis is micropygous, but the pygidium is proportionally larger than that of many other Cambrian trilobites, and there is a slightly variable number of segments in the trunk. Articulation structures are very well developed through the trunk, but interlocking or coaptative devices are poorly developed with the exception of the terrace lines and a novel structure, a ‘posterior arch’, in the pygidium, which would have prevented shear. The absence of genal spines that would otherwise have inhibited sinking into the mud substrate may have enabled Monkaspis daulis to use the enrolment procedure to excavate a hole in the sediment for taking shelter.     

特马豆克期三叶虫Dichelepyge lata Peng的个体发育

记述了产于湘西泸溪特马豆克期锅塘组的刺尾虫类三叶虫Ｄｉｃｈｅｌｅｐｙｇｅｌａｔａ的个体发育过程。其最显著的变化有头鞍形状和比例的改变，头鞍沟、颈沟与背沟的分离．鞍前箍（ｐａｒａｆｒｏｎｔａｌｂａｎｄ）的出现与消失．胸部肋区和轴部比例宽度的改变，尾部的形状和比例的变更，尾边缘的出现和肋沟深度的增加，以及后侧刺的位置逐渐前移。组成成虫期个体的前８个体节（６个胸节和前２个尾节）极有可能在分节０期（ｍｅａｓｐｉｄＤｅｇｒｅｅ０）便已形成。从分节３期（ｍｅｒａｓｐｉｄＤｅｇｒｅｅ３）起在带后侧刺的体节与轴后区（ｐｏｓｔａｘｉａｌｆｉｅｌｄ）之间又发育出一个新的体节。过渡期尾部（ｔｒａｎｓｉｔｏｒｙｐｙｇｉｄｉｕｍ）在分节０期或分节１期发育出后侧刺．在分节４期又同时长出３对侧刺，其中最后一对侧刺与后侧刺一道形成成年期尾部的尾刺。     

Burlingiid trilobites from Norway, with a discussion of their affinities and relationships

Four Middle and Upper Cambrian burlingiid trilobites from the Oslo Region, Norway, are described including Burlingia angusta sp. nov. from the Ptychagnostus punctuosus Zone and Schmalenseeia athrotryphe sp. nov. from the lower part of the Lejopyge laevigata Zone. New complete material previously attributed to Schmalenseeia jagoi Whittington is assigned to Burlingia . Cladistic analysis supports the genera Burlingia and Schmalenseeia as currently understood, including the placement of the controversial middle Middle Cambrian Schmalenseeia acutangula Westergård in Schmalenseeia , even though it lacks typical characters of the genus such as the median ridge on the preglabellar field. The analysis also supports burlingiid monophyly, and suggests that Schmalenseeia was derived from a broadly Burlingia –like ancestor, with S. acutangula displaying how the transition may have occurred. The broader relationships of Burlingia remain obscure, although similarities between burlingiids and the arthropod Kleptothule from the Early Cambrian Sirius Passet fauna are discussed: these include overall form, lack of functional hinges in the thorax, and details of the cephalic region. It is unclear whether these similarities represent general progenetic features, are functional convergences or, less likely, represent a genuine relationship.     

Ontogeny of the Furongian (late Cambrian) trilobite Proceratopyge cf. P. Lata Whitehouse from northern Victoria Land,Antarctica, and the evolution of metamorphosis in trilobites

There were multiple origins of metamorphosis‐undergoing protaspides in trilobite evolution: within the superfamilies Remopleuridioidea, Trinucleoidea, and within the Order Asaphida. Recent studies have revealed that the protaspides of the Cambrian representatives of the Remopleuridioidea and the Trinucleoidea did not undergo metamorphosis. However, ontogeny of the Cambrian members of the Order Asaphida has remained unknown. This study documents the ontogeny of the Furongian asaphoidean ceratopygid trilobite, Proceratopyge cf. P. lata Whitehouse, from northern Victoria Land, Antarctica. Two stages for the protaspid phase, five developmental stages for the post‐protaspid cranidia, and ten stages for the post‐protaspid pygidia have been identified. Interestingly, the protaspis directly developed into a meraspis without metamorphosis. A new cladistic analysis resulted in a single most parsimonious tree, according to which the presence of the bulbous commutavi protaspis turns out to be a synapomorphy for Asaphidae + Cyclopygoidea, not a synapomorphy for the Order Asaphida as previously suggested. In addition, it is inferred that there was convergent evolution of indirectly‐developing commutavi protaspides during the Furongian and Early Ordovician. Metamorphosis‐entailing planktonic larvae evolved in many different metazoan lineages near the Cambrian–Ordovician transition, due to the escalating ecological pressure of the Great Ordovician Biodiversification Event. Since the bulbous commutavi protaspid morphology is thought to be an adaptation for a planktonic life mode, the convergent evolution of the indirect development in the three trilobite lineages at this period might have been a result of adaptation to the early phase of the Great Ordovician Biodiversification Event.     

Two species of Tsinania (Trilobita,Corynexochida) from upper Furongian (Cambrian) of northern Anhui,China and their intraspecific variation

Hundreds of specimens of the trilobite Tsinania were collected from the Cambrian (upper Jiangshanian through Stage 10) at Fenghuangshan in northern Anhui, China, and were identified as Tsinania canens (Walcott, 1905) and Tsinania dolichocephala (Kobayashi, 1933). Comparisons between these two species and others referred to Tsinania, allow the redefinition of the two species. Fourteen synonyms and thirteen synonyms have been incorporated into T. canens and T. dolichocephala respectively. The key diagnosis of T. canens includes: the preglabellar field is flat, relatively short (sag.), has a slightly to obviously angular anterior margin; pygidium is rounded triangular and strongly convex; pygidial border is absent or very narrow. The key diagnosis of Tsinania dolichocephala comprises: the glabella is moderately convex with flat anterior part and convex posterior part (sag.); anterior border is relatively long (sag.), rounded to sharply angulated; pygidium is rounded triangularly with a wide, concave border. Upon such diagnoses, the potential intraspecific variations (e.g., the angularity of the anterior margin, the triangularity of the pygidium, and the width of the pygidial border) of the two species are discussed and listed. The late ontogeny of both species is described to further support the intraspecific variations.     

Thoracic structure and enrolment style in middle Cambrian Eccaparadoxides pradoanus presages caudalization of the derived trilobite trunk

Abstract: The ability to enrol effectively evolved several times among trilobites. Here, we show that, unlike most redlichiid trilobites that could not enrol, both morphotypes of Eccaparadoxides pradoanus from the middle Cambrian of Spain enrolled so as to enclose most of the ventral surface beneath the exoskeleton and possessed specialized articulating devices that facilitated this behaviour. The holaspid thorax of all E. pradoanus was divided into two principal regions. The boundary between these marked a transition from anterior segments with short pleural spines, fulcra and ridge‐and‐groove inner pleural regions to posterior segments with longer, acuminate pleural spines that lack fulcra and inner pleural regions. Devices that aid articulation, such as fulcra with short articulating pleural surfaces, the petaloid articulating facet and long articulating half rings, are concentrated in the anterior region. These features, and the large number of specimens preserved in various degrees of enrolment, suggest an enrolment procedure in which the rear part of the trunk, containing both the posterior thorax and the pygidium, rotated as a single unit without significant internal flexure. As these posterior trunk articulations were apparently not required to permit enrolment, concentrating flexure in the anterior may have presaged the caudalized condition seen in many derived trilobite groups that encapsulated, in which a larger proportion of the trunk segments were allocated to the mature pygidium, and therefore unable to articulate.     

Nematodes associated with Ips cembrae (Coleoptera: Curculionidae): comparison of generations,sexes and sampling methods

A study of nematodes associated with the large larch bark beetle Ips cembrae (Heer 1836) was carried out at three locations in the Czech Republic. The proportion of beetles infested by endoparasitic nematodes (representatives of genera Contortylenchus, Parasitylenchus, Cryptaphelenchus and Parasitorhabditis) ranged from 29.9 to 50.9%. Significant differences were determined in nematode infestation levels among locations, generations and sampling methods. No differences were found in infestation rates between males and females. The percentage of bark beetles with phoretic nematodes ranged from 18 to 42.9%. Phoretic nematodes directly found under elytra, on wings and between body segments of the bark beetles belong to the genus Micoletzkya. However, adults and juveniles of other two phoretic species Laimaphelenchus penardi and Bursaphelenchus sp. were found in the gallery frass of I. cembrae. Infestation by phoretic nematodes positively correlated with the presence of mites under elytra.     

Unravelling the structure and function of the petal appendages in the tribe Schwenckieae (Solanaceae)

• The tribe Schwenckieae (Solanaceae) is characterised by the presence of appendages on the corolla, a diagnostic trait for the group. These appendages constitute a median distal projection of the three‐lobed petal and occur in the genera Melananthus and Schwenckia but are absent in Heteranthia.
• We investigated the micromorphology and anatomical structure of the appendages and lateral petal lobes of Schwenckia americana (two varieties), S. angustifolia, S. curviflora and S. novaveneciana, and Melananthus fasciculatus. We also performed histochemical tests to determine if the appendages are involved in the production of volatiles, acting as a fragrance secretory structure (osmophore).
• The appendages have a uniseriate epidermis, whose cells store phenolics and lipids. The parenchyma is starch‐rich just prior to anthesis in all species studied. The sensory test and anatomical analyses identified scent‐secreting tissues, not only in the appendages, but also in the lateral petal lobes, whose cells are papillose with a sculptured surface. The α‐naphthol p‐phenylenediamine (NADI) reaction detected volatile (essential oils) compounds in S. americana var. americana and S. americana var. angustifolia.
• We demonstrated the secretory tissues and the production of lipids in the corolla appendages of Schwenckia and Melananthus, which indicate their osmogenic function and probable scent emission to attract pollinators.

     

Penetration by Botryosphaeriaceae species in avocado,guava and persimmon fruit during postharvest

Botryosphaeriaceae species have a wide host range and a worldwide distribution. These fungal species can colonize several plant organs, such as the trunk, leaves and fruit. Some Botryosphaeriaceae species cause important diseases on persimmon, avocado and guava fruit. However, there is a lack of information regarding the mechanisms of penetration by Botryosphaeriaceae species on these tropical and subtropical fruits. This study aimed to better understand the mechanisms involved in fungal penetration, host specificity and aggressiveness of Botryosphaeria dothidea, Lasiodiplodia pseudotheobromae and Neofusicoccum parvum on avocado (Persea americana), guava (Psidium guajava) and persimmon (Diospyros kaki) fruit. Scanning electron microscopy (SEM) image analysis showed that in avocado fruit, the three studied Botryosphaeriaceae species penetrated through lenticels. In guava fruit, penetration through stomata was verified for Botryosphaeria dothidea and Neofusicoccum parvum. In persimmon fruit, an appressoria-like structure was observed for B. dothidea, which suggests direct penetration. Disease incidence in wounded fruit was 24% higher than in non-wounded fruit. L. pseudotheobromae and N. parvum showed differences in aggressiveness in guava fruit. The longest incubation period was observed for N. parvum inoculated on guava, with an average of 4.5 days, and the shortest incubation period was verified for B. dothidea inoculated on avocado, with an average of 2.8 days. The area under the disease progress curve (AUDPC) did not differ between Botryosphaeriaceae species on avocado, whereas on guava and persimmon fruit, the AUDPC was lower for B. dothidea. The information regarding penetration mechanisms and aggressiveness is important to improve postharvest disease control strategies.     

Embryogenesis of Heliconius erato (Lepidoptera,Nymphalidae): A contribution to the anatomical development of an evo‐devo model organism

This study reports on the embryogenesis of Heliconius erato phyllis between blastoderm formation and the prehatching larval stage. Syncytial blastoderm formation occurred approximately 2 h after egg laying (AEL) and at about 4 h, the cellular blastoderm was formed. The germ band arose from the entire length of the blastoderm, and rapidly became compacted occupying approximately two‐thirds of the egg length. At about 7 h AEL, protocephalon and protocorm differentiation occurred. Continued proliferation of the germ band was followed by penetration into the yolk mass, forming a C‐shaped embryo at about 10 h. Approximately 12 h AEL, the gnathal, thoracic and abdominal segments became visible. The primordium of the mouthparts and thoracic legs formed as paired evaginations, while the prolegs formed as paired lobes. At about 30 h, the embryo reversed dorsoventrally. Approximately 32 h AEL, the protocephalon and gnathal segments fused, shifting the relative position of the rudimentary appendages in this region. At about 52 h, the embryo was U‐shaped in lateral view and at approximately 56 h, the bristles began evagination from the larval cuticle. Larvae hatched at about 72 h. We found that H. erato phyllis followed an embryonic pattern consistent with long‐germ embryogenesis. Thus, we believe that H. erato phyllis should be classified as a long‐germ lepidopteran. The study of H. erato phyllis embryogenesis provided a structural glimpse into the morphogenetic events that occur in the Heliconius egg period. This study could help future molecular approaches to understanding the evolution of Heliconius development.     

Moulting,ontogeny and sexual dimorphism in the Cambrian ptychopariid trilobite Strenuaeva inflata from the northern Swedish Caledonides

Abstract: Three thousand seven hundred disarticulated remains together with several articulated specimens of the Cambrian Series 2 ptychopariid trilobite Strenuaeva inflata Ahlberg and Bergström, 1978 have been collected from the Torneträsk area, northern Sweden. The material provides significant new data on the morphology, ontogeny, moulting and enrolment of the species. Two distinct morphotypes, possibly an expression of sexual dimorphism, are recognized. The morph with a pair of bulbs in the frontal area, interpreted as brood pouches, is considered to represent females. Statistical treatment of the length/width ratio in cranidia reveals isometric growth during ontogeny for both morphotypes. The transition from the meraspid to holaspid ontogenetic period has been established through recognition of the successive development of the number of thoracic segments in articulated late meraspides. Throughout its life cycle, S. inflata went through 11 meraspid degrees and at least 17 holaspid growth stages. Inferred moult ensembles and exuviae reveal the successive opening of cephalic sutures and the function of the rostral plate during exuviation. As in other ellipsocephalid trilobites in which enrolment is known, the pygidium and two or three thoracic segments of S. inflata are concealed beneath the cephalon (spiral enrolment) during complete enrolment.     

THE LOWER CAMBRIAN EODISCOID TRILOBITE CALODISCUS LOBATUS FROM SWEDEN: MORPHOLOGY, ONTOGENY AND DISTRIBUTION

Abstract:  Several thousand disarticulated remains together with a few complete enrolled specimens of the lower Cambrian eodiscoid trilobite Calodiscus lobatus ( Hall, 1847 ) have been collected at two outcrop areas in Sweden. The material reveals new details of morphology and morphogenesis during ontogenetic development. Size-frequency analyses show that the material from the Fånån rivulet in Jämtland, central Sweden, represents a natural population dominated by juveniles, whereas the material from Gislövshammar in Scania, southern Sweden, has been sorted during postmortem transport. Three stages of protaspid development can be traced and defined as well as all subsequent ontogenetic stages for the cephalon, hypostome and pygidium. The early meraspid pygidium has a pronounced larval notch, which persists, though becoming progressively less distinct in later meraspides. The number of axial rings in the transitory pygidium increases throughout meraspid development until a third and final thoracic segment is liberated. During ontogeny the articulating half-rings are strongly developed, and both meraspides and holaspides were capable of full sphaeroidal enrollment and outstretched postures. The hypostome undergoes some dramatic modifications; in M0 the anterior margin is axe-shaped, by M1 the area of attachment greatly decreases and the hypostome becomes more elongated and pear-shaped, before attaining its adult form, which has an overall resemblance to that of polymerid trilobites. During ontogeny, the hypostome changes from a conterminant attachment to a natant condition, thereby mirroring hypostomal evolution within trilobites generally. The morphology, ontogeny, enrollment, hypostomal development and the presence of calcified protaspides suggest polymerid rather than agnostoid affinities of the eodiscoids.     

Stink bug rostrum length vs. stylet penetration potential

Stink bugs (Hemiptera: Pentatomidae) and related species continue to plague cotton, Gossypium hirsutum L. (Malvaceae), worldwide. Stink bugs utilize their stylets (housed within the rostrum) to feed upon cotton bolls and transmit pathogens that cause seed and boll rot of cotton. Stylet penetration potential of stink bugs is influenced by species and recent observations indicated a phenomenon whereby stink bugs with shorter rostra yielded deeper stylet penetration estimates. The objective of this study was to elucidate the relationship between rostrum length and known stylet penetration estimates for two pairs of similar‐sized pentatomid species: Chinavia hilaris (Say) vs. Euschistus servus (Say), and Oebalus pugnax (Fabricius) vs. Piezodorus guildinii Westwood. For each species, individual rostral segments were measured to yield total lengths, and measurements were compared against known stylet penetration estimates. Chinavia hilaris and P. guildinii have longer rostra than E. servus and O. pugnax, respectively, yet E. servus and O. pugnax yielded deeper stylet penetration estimates. Deeper stylet penetration by species with shorter rostra can be attributed to differences in the lengths of rostral segments 1 and 2. Euschistus servus and O. pugnax each had significantly longer rostral segments 1 and 2 than C. hilaris and P. guildinii, respectively. Also, the cumulative lengths of rostral segments 1 and 2 comprised a higher overall proportion of the entire rostrum length in E. servus and O. pugnax vs. C. hilaris and P. guildinii, respectively. Rostral segments 1 and 2 are instrumental in the feeding mechanics of these phytophagous species; it is clear that their greater length and their role in stylet penetration model calculations – including the lengths of segments 3 and 4 – override the presumption that total rostrum length equates to stylet penetration potential. This novel finding contributes to the general knowledge of stink bug feeding mechanics.     

Cooperation between the chloroplast psbA 5′‐untranslated region and coding region is important for translational initiation: the chloroplast translation machinery cannot read a human viral gene coding region

Chloroplast mRNA translation is regulated by the 5′‐untranslated region (5′‐UTR). Chloroplast 5′‐UTRs also support translation of the coding regions of heterologous genes. Using an in vitro translation system from tobacco chloroplasts, we detected no translation from a human immunodeficiency virus tat coding region fused directly to the tobacco chloroplast psbA 5′‐UTR. This lack of apparent translation could have been due to rapid degradation of mRNA templates or synthesized protein products. Replacing the psbA 5′‐UTR with the E. coli phage T7 gene 10 5′‐UTR, a highly active 5′‐UTR, and substituting synonymous codons led to some translation of the tat coding region. The Tat protein thus synthesized was stable during translation reactions. No significant degradation of the added tat mRNAs was observed after translation reactions. These results excluded the above two possibilities and confirmed that the tat coding region prevented its own translation. The tat coding region was then fused to the psbA 5′‐UTR with a cognate 5′‐coding segment. Significant translation was detected from the tat coding region when fused after 10 or more codons. That is, translation could be initiated from the tat coding region once translation had started, indicating that the tat coding region inhibits translational initiation but not elongation. Hence, cooperation/compatibility between the 5′‐UTR and its coding region is important for translational initiation.     

Dysregulated molecular pathways in amyotrophic lateral sclerosis–frontotemporal dementia spectrum disorder

Frontotemporal dementia (FTD), the second most common form of dementia in people under 65 years of age, is characterized by progressive atrophy of the frontal and/or temporal lobes. FTD overlaps extensively with the motor neuron disease amyotrophic lateral sclerosis (ALS), especially at the genetic level. Both FTD and ALS can be caused by many mutations in the same set of genes; the most prevalent of these mutations is a GGGGCC repeat expansion in the first intron of C9ORF72. As shown by recent intensive studies, some key cellular pathways are dysregulated in the ALS‐FTD spectrum disorder, including autophagy, nucleocytoplasmic transport, DNA damage repair, pre‐mRNA splicing, stress granule dynamics, and others. These exciting advances reveal the complexity of the pathogenic mechanisms of FTD and ALS and suggest promising molecular targets for future therapeutic interventions in these devastating disorders.     

A genomic assessment of species boundaries and hybridization in a group of highly polymorphic anoles (distichus species complex)

Delimiting young species is one of the great challenges of systematic biology, particularly when the species in question exhibit little morphological divergence. Anolis distichus, a trunk anole with more than a dozen subspecies that are defined primarily by dewlap color, may actually represent several independent evolutionary lineages. To test this, we utilized amplified fragment length polymorphisms (AFLP) genome scans and genetic clustering analyses in conjunction with a coalescent‐based species delimitation method. We examined a geographically widespread set of samples and two heavily sampled hybrid zones. We find that genetic divergence is associated with a major biogeographic barrier, the Hispaniolan paleo‐island boundary, but not with dewlap color. Additionally, we find support for hypotheses regarding colonization of two Hispaniolan satellite islands and the Bahamas from mainland Hispaniola. Our results show that A. distichus is composed of seven distinct evolutionary lineages still experiencing a limited degree of gene flow. We suggest that A. distichus merits taxonomic revision, but that dewlap color cannot be relied upon as the primary diagnostic character.