Ontogeny of the redlichiid trilobite Eoredlichia intermediata from the Chengjiang Lagerstätte,lower Cambrian,southwest China

A detailed investigation of the morphology and ontogeny of the redlichioid trilobite Eoredlichia intermediata (Bulletin of the Geological Society of China, 3–4, 1940, 333) from the lower Cambrian Yu'anshan Member of the Heilinpu Formation, in Kunming, Yunnan Province, southwest China, is presented. The new material comprises a relatively complete ontogenetic series ranging from the early meraspid to the holaspid period, which reveals more details on morphological variation such as the appearance of bacculae in some holaspid specimens, contraction and disappearance of fixigenal spines, and macropleural spines of the first and second thoracic segments, which are all documented for the first time and can also be used as developmental markers defining ontogenetic phases. Two distinct morphotypes, possibly an expression of intraspecific variation or sexual dimorphism, are distinguished by the morphology of pleural spines of the second thoracic segment in meraspid degree 14 and holaspides. The trunk segmentation schedule of E. intermediata is also discussed and conforms to the protarthrous developmental mode. The distinction of the thoracic region into two portions can be observed during late meraspid and early holaspid periods, which might be considered as a reference for a better understanding on the relationship of tagmosis and growth segmentation among the Cambrian redlichiid trilobites.     

Trilobite body patterning and the evolution of arthropod tagmosis

Preservation permitting patterns of developmental evolution can be reconstructed within long extinct clades, and the rich fossil record of trilobite ontogeny and phylogeny provides an unparalleled opportunity for doing so. Furthermore, knowledge of Hox gene expression patterns among living arthropods permit inferences about possible Hox gene deployment in trilobites. The trilobite anteroposterior body plan is consistent with recent suggestions that basal euarthropods had a relatively low degree of tagmosis among cephalic limbs, possibly related to overlapping expression domains of cephalic Hox genes. Trilobite trunk segments appeared sequentially at a subterminal generative zone, and were exchanged between regions of fused and freely articulating segments during growth. Homonomous trunk segment shape and gradual size transition were apparently phylogenetically basal conditions and suggest a single trunk tagma. Several derived clades independently evolved functionally distinct tagmata within the trunk, apparently exchanging flexible segment numbers for greater regionally autonomy. The trilobite trunk chronicles how different aspects of arthropod segmentation coevolved as the degree of tagmosis increased.     

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.     

Pair-rule genes cooperate to activate en stripe 15 and refine its margins during germ band elongation in the D. melanogaster embryo

Patterns of gene expression have been well documented during embryogenesis for the Drosophila melanogaster trunk segments. The same is not the case for the terminal segments. Here, gene expression patterns are followed during embryogenesis in the caudal segments (A8-A10 and the anal plate), with special attention paid to the novel regulation of engrailed (en). Chosen for this study are the pair-rule genes even-skipped (eve), fushi tarazu (ftz), runt (run), hairy (h), paired (prd) and odd-skipped (odd), and the segment polarity gene (en). The results demonstrate a progressive and coupled translocation of gene expression distally for all genes studied, suggesting that the most posterior segments are determined later than trunk segments.     

Positional specification in the segmental growth pattern of an early arthropod

In many arthropods, there is a change in relative segment size during post-embryonic development, but how segment differential growth is produced is little known. A new dataset of the highest quality specimens of the 429 Myr old trilobite Aulacopleura koninckii provides an unparalleled opportunity to investigate segment growth dynamics and its control in an early arthropod. Morphometric analysis across nine post-embryonic stages revealed a growth gradient in the trunk of A. koninckii. We contrastively tested different growth models referable to two distinct hypotheses of growth control for the developing trunk: (i) a segment-specific control, with individual segments having differential autonomous growth progression, and (ii) a regional control, with segment growth depending on their relative position along the main axis. We show that the trunk growth pattern of A. koninckii was consistent with a regional growth control producing a continuous growth gradient that was stable across all developmental stages investigated. The specific posterior-to-anterior decaying shape of the growth gradient suggests it deriving from the linear transduction of a graded signal, similar to those commonly provided by morphogens. A growth control depending on a form of positional specification, possibly realized through the linear interpretation of a graded signal, may represent the primitive condition for arthropod differential growth along the main body axis, from which the diverse and generally more complex forms of growth control in subsequent arthropods have evolved.     

Larval development of Japanese 'conchostracans': part 2, larval development of Caenestheriella gifuensis (Crustacea, Branchiopoda, Spinicaudata, Cyzicidae), with notes on homologies and evolution of certain naupliar appendages within the Branchiopoda

As part of a larger project examining and comparing the ontogeny of all major taxa of the Branchiopoda in a phylogenetic context, the larval development of Caenestheriella gifuensis (Ishikawa, 1895), a Japanese spinicaudatan ‘conchostracan’, is described by scanning electron microscopy. Seven different larval stages are recognised, in most cases based on significant morphological differences. They range in length from about 200 to 850 μm. Nauplius 1 has a plumb and lecithotrophic appearance with a rounded hind body and a labrum with an incipient medial spine. Limb segmentation is mostly unclear but the second antennae have more putative segments delineated than are expressed in the later stages. Feeding structures such as the mandibular coxal process and antennal coxal spine are only weakly developed. Nauplius 2 is very different from nauplius 1 and has three large spines on the labral margin and two long caudal spines. Feeding structures such as the mandibular coxal process and various spines and setae are developed, but whether feeding begins at this stage was not determined. The mandible has developed an ‘extra’ seta on endopod segment 1, absent in Nauplius 1. The segmentation of the second antenna has changed significantly due to fusions of various early segments. Nauplius 3 is like nauplius 2 in morphological detail, but larger and more elongate. Nauplius 4 has developed a pair of small anlagen of the carapace and rudiments of the first five pairs of trunk limbs, and the coxal spine of the antenna has become distally bifid. Nauplius 5 has a larger carapace anlage, externally visible enditic portions of the elongate trunk limbs, and a pair of primordial dorsal telson setae. Nauplius 6 has a larger and partly free carapace and better-developed, partly free trunk limbs with incipient enditic, endopodal, and exopodal setation. A pair of caudal spines, dorsal to the large caudal spines, has appeared. Nauplius 7 is quite similar to nauplius 6 but is larger and has slightly longer caudal and labral spines; also, the setation of the most anterior trunks limbs is better developed. The larval development is largely similar to that of other spinicaudatans. The larval mandible, which is evolutionarily conservative within the Branchiopoda, reveals a setation pattern similar to that of the Anostraca and Notostraca (two setae on mandibular endopod segment 1). Most other spinicaudatans and all examined laevicaudatans share another setal pattern (one seta on mandibular endopod segment 1), which could indicate a close relationship among these taxa. The second antenna undergoes a special development, which provides an insight into the evolution of this limb within the Branchiopoda. In nauplius 1 the basipod, endopod, and exopod are all superficially divided into a relatively high number of segments. In later nauplii some of these have fused, forming fewer but larger segments. We suggest that this ontogeny reflects the evolution of antennae in the conchostracans. Various aspects of the morphology of the antennae are discussed as possible synapormorphies for either the Diplostraca or subgroups of the Conchostraca.     

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.     

<Emphasis Type="Italic">Tubulideres seminoli</Emphasis> gen. et sp. nov. and <Emphasis Type="Italic">Zelinkaderes brightae</Emphasis> sp. nov. (Kinorhyncha,Cyclorhagida) from Florida

One new kinorhynch genus and species and one new species from the genus Zelinkaderes are described from sandy sediment off Fort Pierce, Florida. The new genus and species, Tubulideres seminoli gen. et sp. nov. is characterized by the presence of the first trunk segment consisting of a closed ring, the second segment of a bent tergal plate with a midventral articulation and the following nine segments consisting of a tergal and two sternal plates. Cuspidate spines are not present, but flexible tubules are located on several segments, and in particular concentrated on the ventral side of the second segment. Middorsal spines are present on all trunk segments and are alternatingly offset to a position slightly lateral to the middorsal line. Zelinkaderes brightae nov. sp. is characterized by its spine formula in having middorsal spines on trunk segments 4, 6 and 8–11, lateroventral acicular spines on segment 2, lateral accessory cuspidate spines on segments 2 and 8, ventrolateral cuspidate spines on segments 4–6 and 9, lateroventral acicular spines present on segments 8 and 9, and midterminal, lateral terminal and lateral terminal accessory spines on segment 11. The spine formula of Z. brightae nov. sp. places it in a position in between Z. submersus and a clade consisting of Z. klepali and Z. floridensis. The new findings on Z. brightae nov. sp. have led us to propose an emended diagnosis for the genus.     

Structure, ontogeny, and moulting of the olenid trilobite Ctenopyge (Eoctenopyge) angusta Westergård, 1922 from the upper Cambrian of Västergötland, Sweden

The genus Ctenopyge is known mainly from disarticulated sclerites and from rare complete specimens flattened in shales. Hitherto, very few specimens have been found preserved intact and in three dimensions. In a recently discovered fauna, however, in the Peltura minor Subzone in Västergötland, central Sweden, there occur several species of Ctenopyge , of which many are complete and superbly preserved; moreover they occur at all stages of growth. Of these the abundant Ctenopyge ( Eoctenopyge ) angusta Westergård, 1922 is described and reconstructed here as an adult, and the entire ontogeny is documented for all post–protaspid growth stages. Many characters typical of the adult, such as the long genal spines and the caudal spine, develop very early in ontogeny, and the relative dimensions of the cranidium do not greatly change during growth. Macropleural spines, however, develop later. The transitory pygidium, relatively large and shield–shaped in the early meraspid, later becomes very small as the ten thoracic segments are liberated; a median spine develops on the last thoracic segment only at the holaspid stage. Instar groupings can be clearly distinguished for the early stages. Recurrent associations of sclerites are interpreted as moulting configurations. As reconstructed, the genal spines are horizontal and parallel with the extended thorax; an adaptation which presumably allowed the trilobite to rest on the sea floor.     

Trunk segmentation of Cambrian eodiscoid trilobites

SUMMARY Here we report, from Cambrian Series 2 of Chongqing, southern China, on some three-dimensionally phosphatized exoskeletons representing a series of instars of a specialized eodiscoid trilobite Badiscus spinosus gen. et sp. nov. The preservation is unusual and its protaspides (the youngest juveniles) belonging to different growth stages markedly exhibit some "embryonic trunk segments," which imply the specification of all body segments during embryogenesis. This previously unknown style of tagmosis, as being inconsistent with the conventional view of sequential segmentation during postembryonic anamorphosis, emphasizes the complexity of trilobite trunk segmentation during ontogeny and indicates for the first time that epimorphic development, like other known patterns during postembryonic development, may have already been adopted by some of the earliest trilobites.     

Growth and variation in the Silurian proetide trilobite Aulacopleura konincki and its implications for trilobite palaeobiology

Morphometric analysis of growth in Aulacopleura konincki reveals several important features: (1) morphological variability is approximately constant throughout growth, increasing slightly in later ontogeny; (2) shape is more tightly constrained than the numbers of postcephalic segments, which can be quite variable; and (3) a major ontogenetic transition occurs at glabellar lengths of about 1.7 mm. This transition divides the ontogeny into two distinct growth phases, is smooth rather than abrupt, and is expressed as changes in growth trajectories, especially in the pygidium. The transition is not strictly correlated with the number of thoracic or pygidial segments. These results suggest a re-evaluation of the concepts of meraspid and holaspid growth stages in A. konincki , using growth trajectories rather than thoracic segment number to define the stages. Developmental flexibility in holaspid segment numbers in this phylogenetically advanced trilobite suggests that environmental factors, rather than phylogenetic constraint, may explain variations in segment production patterns within some advanced Trilobita. Morphometrics, trilobites, ontogeny, variability, segmentation, Silurian, Czech Republic.     

Ancestral Patterning of Tergite Formation in a Centipede Suggests Derived Mode of Trunk Segmentation in Trilobites

Trilobites have a rich and abundant fossil record, but little is known about the intrinsic mechanisms that orchestrate their body organization. To date, there is disagreement regarding the correspondence, or lack thereof, of the segmental units that constitute the trilobite trunk and their associated exoskeletal elements. The phylogenetic position of trilobites within total-group Euarthropoda, however, allows inferences about the underlying organization in these extinct taxa to be made, as some of the fundamental genetic processes for constructing the trunk segments are remarkably conserved among living arthropods. One example is the expression of the segment polarity gene engrailed, which at embryonic and early postembryonic stages is expressed in extant panarthropods (i.e. tardigrades, onychophorans, euarthropods) as transverse stripes that define the posteriormost region of each trunk segment. Due to its conservative morphology and allegedly primitive trunk tagmosis, we have utilized the centipede Strigamia maritima to study the correspondence between the expression of engrailed during late embryonic to postembryonic stages, and the development of the dorsal exoskeletal plates (i.e. tergites). The results corroborate the close correlation between the formation of the tergite borders and the dorsal expression of engrailed, and suggest that this association represents a symplesiomorphy within Euarthropoda. This correspondence between the genetic and phenetic levels enables making accurate inferences about the dorsoventral expression domains of engrailed in the trunk of exceptionally preserved trilobites and their close relatives, and is suggestive of the widespread occurrence of a distinct type of genetic segmental mismatch in these extinct arthropods. The metameric organization of the digestive tract in trilobites provides further support to this new interpretation. The wider evolutionary implications of these findings suggest the presence of a derived morphogenetic patterning mechanism responsible for the reiterated occurrence of different types of trunk dorsoventral segmental mismatch in several phylogenetically distant, extinct and extant, arthropod groups.     

Postembryonic development of Antygomonas incomitata (Kinorhyncha: Cyclorhagida)

Postembryonic development in the kinorhynch species Antygomonas incomitata was examined using scanning electron microscopy. The morphology of the six juvenile stages, J‐1 to J‐6, varies at numerous details, but they can also be distinguished by a few key characters. Juvenile stage 1 by its composition of only nine trunk segments; J‐2 by the combination of possessing 10 trunk segments, but no cuspidate spines on segment 9; J‐3 by the presence of cuspidate spines on segment 9, but only one pair of cuspidate spines on segment 8; J‐4 by the combination of 10 trunk segments only, but having two pairs of cuspidate spines on segment 8; J‐5 by possessing 11 trunk segments and same spine compositions as adults but is still maintaining postmarginal spiculae; J‐6 specimens closely resemble adults and are most easily identified by their reduced trunk lengths. New segments are formed in a growth zone in the anterior part of the terminal segment. The complete number of segments is reached in J‐5. Development of cuticular head and trunk structures are described through all postembryonic stages and following developmental patterns could be outlined: the mouth cone possesses outer oral styles from J‐1, but in J‐1 to J‐3, the styles alternate in size. Scalids of the introvert are added after each molt, and scalids appear earliest in the anterior rings, whereas scalids in more posterior rings are added in older postembryonic stages. The early J‐1 stage is poor in spines and sensory spots and both structures increase in number after each molt. The complete spine composition is reached in J‐4, whereas new sensory spots appear after all molts, inclusive the final one from J‐6 to adult. Sensory spots in the paraventral positions often appear as Type 3 sensory spots but are through development transformed to Type 2. This transformation happens earliest on the anterior segments. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

Two new species in the Echinoderes coulli group (Echinoderidae,Cyclorhagida, Kinorhyncha) from the Ryukyu Islands,Japan

Two new species belonging to the Echinoderes coulli group are described with their external morphologies and sequences of nuclear 18S rRNA and 28S rRNA genes, and mitochondrial COI gene. The first species, Echinoderes komatsui sp. n., is characterized by absence of acicular spines, and presence of lateroventral tubules on segments 5 and 8, laterodorsal tubules on segment 10, inverted triangle or wide oval shaped large sieve plates, lateral terminal accessory spines in female, and short tips of ventral pectinate fringe on segment 10. The second species, Echinoderes hwiizaa sp. n., is characterized by absence of acicular spines, and presence of lateroventral tubules on segments 5 and 7–9, midlateral tubules on segment 8, laterodorsal tubules on segment 10, large narrow oval shaped sieve plates on segment 9, and thick, short and blunt lateral terminal spines about 10–15% of trunk length. The diagnostic characters and key to species of E. coulli group are provided as well.     

Kinorhyncha from the stomach of the shrimp Pleoticus muelleri (Bate, 1888) from Comodoro Rivadavia, Argentina

Six kinorhynchs were found in the stomachs of the Argentine red shrimp, Pleoticus mulleri (Bate, 1888) from the Argentine coast of Patagonia. Three new species are described: Condyloderes storchi n. sp., Pycnophyes argentinensis n. sp. and P. neuhausi n. sp. A fourth species, Kinorhynchus anomalus Lang, 1953 was previously known only from the coast of Chile. This is the third known record of kinorhynchs documented as a food source. Condyloderes storchi, n. sp. is the fourth new species in this genus. It is distinguished by its paradorsal cuspidate spines on segments 7 and 9, lateral accessory and ventrolateral spines on segments 3, 6, 7, 10 and 11. P. argentinensis, n. sp. has nearly equal sternal width for segments 3–11 (about 7% of the trunk length), episternal plates with three distinct areas along the anterior margin, mid-sternal plate with even margin, mid-dorsal spinose protrusions along the terminal borders of segments 11 and 12, and lateral terminal spines 176 μm long, about 21% of trunk length. P. neuhausi, n. sp.has a prominent posterior elongation of the tergal plate of segment 3, uneven lateral margins of the mid-sternal plate, a maximum sternal width at segment 3, no mid-dorsal spinose processes and mid-ventral thickenings on segments 10–12.     

Enrolment and trunk segmentation of a Cambrian eodiscoid trilobite

Completely enrolled, phosphatized, 3‐D specimens of the eodiscinid trilobite Tsunyidiscus yanjiazhaiensis from Cambrian Stage 3 of South China exhibit much morphological detail and show variation in delicate coaptative structures associated with their trunk segmentation. The relationship between enrolment mechanism and trunk segmentation during ontogeny confirms a unique developmental pattern among the early Cambrian eodiscoid trilobites, revealing how these animals controlled the rate of segment increase and release during post‐embryonic development in their life cycles.     

Morphology and ultrastructure of the Malpighian tubules of the Chilean common tarantula (Araneae: Theraphosidae).

Relatively little is known about the morphology and ultrastructure of the Malpighian tubules of spiders (Arachnida: Araneae). Our study represents the first investigation of the Malpighian tubules of a theraphosid spider and is the only study to examine the living Malpighian tubules using confocal laser scanning microscopy. In theraphosid spiders, the Malpighian tubules originate from the stercoral pocket in the posterior portion of the opisthosoma and extend forward toward the prosoma in a dendritic pattern. There are three distinct segments (initial, main, and terminal), all dark brown in appearance. Each segment has distinctive ultrastructural features. Both the terminal and the main segment appear to be composed of at least two cell types with finger-like cytoplasmic protrusions associated with one of these types. The terminal segment, which is most proximal to the stercoral pocket, is the largest in diameter. It is composed of large, cuboidal cells containing many mitochondria and lipid inclusions. The main segment is intermediate in diameter with many mitochondria and secretory vesicles present. The initial segment is relatively thin in comparison to the other segments and is intimately associated with the digestive gland. The cells of the initial segment contain very little cytoplasm, fewer mitochondria, secretory vesicles, and prominent inclusions.     

A morphometric analysis of the spinal motor pool in relation to its target muscle during growth in the trout, Salmo gairdneri Richardson

The relation between number and size of spinal motoneurons and the dimension of myotomal muscle has been investigated in trout at different stages of embryonic, larval and postlarval development (body length 1–15 cm). Three spinal segments have been analysed (cervical, trunk and caudal) and the following parameters were determined by means of a Micromeasurements Image Analyzer: (a) mean cross-sectional myotomal area; (b) mean soma size of principal (or dorsomedian, DM) and secondary (or ventrolateral, VL) motoneurons; (c) DM and VL motoneuron density per segment. Myotomal muscle and motor pool growth was evaluated by percent increments of a, b and c parameters at each stage. Their relationships were denned by equations of computed regression lines.
The analysis provided evidence that: (1) a continuous exponential growth of mean myotomal area takes place in the three segments, with the same trend and with the lowest values in the caudal segment; (2) DM and VL motoneuron size and density per segment also increase during development, with the least value in the caudal segment, VL parameters being of lesser value than DM; (3) motoneuron pool and its target myotomal muscle parameters bear a linear relationship as defined by equations of computer regression lines; (4) motoneuron number percent increment at early eleutherembryonic stage precedes myotomal area increment which takes place during late eleutherembryonic stage.
It is apparent that spinal motor pool and target myotomal muscle grow at the same rate in the trout during the considered stages. The discussion links this fact with the hypothesis of a neuronal influence on muscle fibre type differentiation.     

Trilobite tagmosis and body patterning from morphological and developmental perspectives

The Trilobita were characterized by a cephalic region in whichthe biomineralized exoskeleton showed relatively high morphologicaldifferentiation among a taxonomically stable set of well definedsegments, and an ontogenetically and taxonomically dynamic trunkregion in which both exoskeletal segments and ventral appendageswere similar in overall form. Ventral appendages were homonomousbiramous limbs throughout both the cephalon and trunk, exceptfor the most anterior appendage pair that was antenniform, preoral,and uniramous, and a posteriormost pair of antenniform cerci,known only in one species. In some clades trunk exoskeletalsegments were divided into two batches. In some, but not all,of these clades the boundary between batches coincided withthe boundary between the thorax and the adult pygidium. Therepeated differentiation of the trunk into two batches of segmentsfrom the homonomous trunk condition indicates an evolutionarytrend in aspects of body patterning regulation that was achievedindependently in several trilobite clades. The phylogeneticplacement of trilobites and congruence of broad patterns oftagmosis with those seen among extant arthropods suggest thatthe expression domains of trilobite cephalic Hox genes may haveoverlapped in a manner similar to that seen among extant arachnates.This, coupled with the fact that trilobites likely possessedten Hox genes, presents one alternative to a recent model inwhich Hox gene distribution in trilobites was equated to eightputative divisions of the trilobite body plan.