ALBERT

Description: Cambrian biostratigraphy of the Indian subcontinent is best documented from the Parahio Formation of the Tethyan Himalaya. Recently established trilobite biostratigraphy shows that the formation encompasses the latest part of unnamed Stage 4 and much of unnamed Stage 5. A variety of small shelly fossils have been recovered via acid digestion of carbonate beds and include tetract and pentact hexactinellid sponge spicules, chancelloriid spicules belonging to Chancelloria sp. and a new species, Archiasterella dhiraji , shells of an helcionelloid comparable to Igorella maidipingensis , a meraspid ptychopariid trilobite, the tubular Cupitheca sp., a poorly preserved hyolith, and an assortment of spinose microfossils of uncertain affinity. These newly recovered microfossils are consistent with the trilobite-based lower and middle Cambrian age determination and do not support a late Cambrian age for the top of the Parahio Formation advocated in some recent literature. The microfossils reported herein significantly expand the known diversity of such fossils from Cambrian strata in the Himalayan region, and allow for comparison of this fauna with others from Gondwanaland and elsewhere. Integration with trilobite data indicate that the stratigraphic ranges of many small shelly fossils described in this study are greater than previously recognized.

Description: A bstract A new dataset of the highest quality specimens of fully articulated, juvenile and mature exoskeletons of the Czech middle Silurian trilobite Aulacopleura koninckii offers improved resolution of original morphology by all measures considered. The degree of variation in both size and shape among later meraspid instars was constant, and suggesting targeted growth in both attributes. Size-related changes in the shape of the dorsal exoskeleton and of the segment-invariant cephalon were detected in the meraspid stage, but in the holaspid phase marked allometry was detected only in the trunk region, with the pygidium showing notable expansion in relative size. Meraspid cranidial allometry was subtle, with significant changes in instar form detectable only after several molts. This trilobite developed gradually throughout meraspid and holaspid ontogeny, with the synchronous cessation of trunk segment appearance and release at the onset of the holaspid phase. Precise development of shape and size occurs in the context of marked variability in the number of trunk segments at maturity, illustrating complex patterns of character variation within a species. A new systematic description establishes the synonymy of several subspecies with A. koninckii .

Description: New discoveries of the early Cambrian yiliangellinine trilobite Zhangshania typica Li and Zhang in Kunming preserve almost all instars from early postembryonic (protaspid) to mature (holaspid) phases in articulated state, in addition to mature specimens with antennae bearing paired spines on the basal articles. The ontogenetic series shows protarthrous development with some, but likely not all, early holaspid instars expressing additional pygidial segments, gradual rearward migration of the location of the longest pleural spines on the trunk segments, and striking positive allometry of the genal spines. It also reveals Parazhangshania sichuanensis Li and Zhang, 1990 to be the holaspid stage 3 of Z . typica, and therefore its junior synonym. This new find in the Hongjingshao Formation provides species-based regional correlation across the South China block and Z . typica may provide an important biostratigraphic marker for the base of the traditional Tsanglangpuan Stage.

Description: A bstract Onset of maturity in trilobites is generally considered to occur when the last trunk segment is released into the thorax, marking the start of the holaspid stage. Here we describe striking morphological changes that occur within the holaspid ontogeny of Lonchopygella megaspina Zhou in Zhou et al., 1977 , which include the effacement of dorsal furrows, the rapid and complete degeneration of pygidial lateral spines, and the increasing prominence of a pygidial axial spine. These notable changes, which are not coincident with the onset of the holaspid phase, emphasize that the onset of maturity in trilobites should be viewed on a character-by-character basis before assessing whether the exoskeleton as a whole can be described as mature. The holaspid pygidial condition in L. megaspina may represent an intermediate step in an evolutionary transition in the number, form, and allocation of segments in the tsinaniid trunk. Pygidial transition from a dynamically changing complement of segments in the meraspid phase to a static complement in the holaspid phase was accompanied by a marked change in the extent to which segment boundaries defined pygidial structure. Attaining this static complement allowed subsequent pygidial development to emphasize its structure as an integrated unit in which internal segmental boundaries became diffuse, a continuous margin to become prominent, and an elongated terminal spine, first evident at onset of epimorphic growth, to develop allometrically. Trilobite body development suggests that while the segmented construction placed constraints on how morphology varied, the influence of these constraints diminished following completion of thoracic segment construction. Selective premium for a distinct posterior tagma might favor the early ontogenetic acquisition of such a structure, and could have been a driver of the repeated trend toward caudalization witnessed among derived trilobite clades.

Description: Recent morphometric analysis revealed a juvenile (meraspid) axial growth gradient in the trunk of the ~429 Myr old trilobite Aulacopleura koninckii that resulted from growth control based on positional specification, as is common among extant organisms. Here we explore axial growth gradients in the more anterior body region, the cephalon, and in the cephalon and trunk during subsequent development in the holaspid period. We detected an axial growth gradient in the cephalon in the meraspid period, flatter and opposite in direction to that of the trunk, which also persisted during the holaspid period. We also found an holaspid trunk growth gradient, with a different distribution of growth rates among segments than that of the meraspid period. These newly observed growth gradients are compatible with the mechanism of growth control inferred for the meraspid trunk. Thus, the same kind of growth control may have operated in both body regions and during the whole ontogeny of A. koninckii. This study, along with others on the same species that preceded it, show that morphometric analysis of appropriate data sets can address questions of high interest for evolutionary developmental biology using data from fossils. By revealing developmental features at deep nodes of the phylogenetic tree, these studies will elucidate both how developmental processes evolved and how they themselves affected the evolution of organismal body patterning.