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result(s) for
"Topper, Timothy P."
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An encrusting kleptoparasite-host interaction from the early Cambrian
2020
Parasite–host systems are pervasive in nature but are extremely difficult to convincingly identify in the fossil record. Here we report quantitative evidence of parasitism in the form of a unique, enduring life association between tube-dwelling organisms encrusted to densely clustered shells of a monospecific organophosphatic brachiopod assemblage from the lower Cambrian (Stage 4) of South China. Brachiopods with encrusting tubes have decreased biomass (indicating reduced fitness) compared to individuals without tubes. The encrusting tubes orient tightly in vectors matching the laminar feeding currents of the host, suggesting kleptoparasitism. With no convincing parasite–host interactions known from the Ediacaran, this widespread sessile association reveals intimate parasite–host animal systems arose in early Cambrian benthic communities and their emergence may have played a key role in driving the evolutionary and ecological innovations associated with the Cambrian radiation.
Parasitic interactions are difficult to document in the fossil record. Here, Zhang et al. analyze a large population of a Cambrian brachiopod and show it was frequently encrusted by tubes aligned to its feeding currents and that encrustation was associated with reduced biomass, suggesting a fitness cost.
Journal Article
A stem group echinoderm from the basal Cambrian of China and the origins of Ambulacraria
by
Skovsted, Christian B.
,
Topper, Timothy P.
,
Guo, Junfeng
in
147/135
,
631/181/2480
,
631/181/414
2019
Deuterostomes are a morphologically disparate clade, encompassing the chordates (including vertebrates), the hemichordates (the vermiform enteropneusts and the colonial tube-dwelling pterobranchs) and the echinoderms (including starfish). Although deuterostomes are considered monophyletic, the inter-relationships between the three clades remain highly contentious. Here we report,
Yanjiahella biscarpa
, a bilaterally symmetrical, solitary metazoan from the early Cambrian (Fortunian) of China with a characteristic echinoderm-like plated theca, a muscular stalk reminiscent of the hemichordates and a pair of feeding appendages. Our phylogenetic analysis indicates that
Y
.
biscarpa
is a stem-echinoderm and not only is this species the oldest and most basal echinoderm, but it also predates all known hemichordates, and is among the earliest deuterostomes. This taxon confirms that echinoderms acquired plating before pentaradial symmetry and that their history is rooted in bilateral forms.
Yanjiahella biscarpa
shares morphological similarities with both enteropneusts and echinoderms, indicating that the enteropneust body plan is ancestral within hemichordates.
The early evolution of the deuterostomes is not well resolved. Here, Topper and colleagues investigate the early Cambrian metazoan
Yanjiahella biscarpa
, concluding that it is a stem echinoderm, is among the oldest known deuterstomes, and supports an ancestral enteropneust body plan in hemichordates.
Journal Article
Symbiotic interactions on middle Cambrian echinoderms reveal the oldest parasitism on deuterostomes
2025
Biotic interactions are considered as major drivers of evolutionary changes, but their identification in the fossil record is extremely rare and controversial. Based on qualitative and quantitative analyses, we report evidence of a biotic interaction between an echinoderm host and its symbiont, probably a parasitic epibiont, from the Cambrian Wuliuan Stage of Australia. The echinoderm plates bear external outgrowths with a median pit at their distal end. These unusual structures have not been previously documented from Cambrian echinoderms and their lack of consistency across various morphological parameters, supports the interpretation that a biotic interaction generated these unique structures. Perturbations in plate microstructure and the overproduction of skeletal material in specific regions, together with reduced size, negatively impact the host’s growth suggesting a parasitic interaction. This reaction by the echinoderm host may represent the progressive embedment of the invasive epibiont. This record represents the oldest evidence of parasitism among deuterostomes.
Journal Article
Evolution and diversity of biomineralized columnar architecture in early Cambrian phosphatic-shelled brachiopods
by
Zhang, Zhiliang
,
Pan, Bing
,
Topper, Timothy P
in
Animals
,
Architecture
,
biologically controlled
2024
Biologically-controlled mineralization producing organic-inorganic composites (hard skeletons) by metazoan biomineralizers has been an evolutionary innovation since the earliest Cambrian. Among them, linguliform brachiopods are one of the key invertebrates that secrete calcium phosphate minerals to build their shells. One of the most distinct shell structures is the organo-phosphatic cylindrical column exclusive to phosphatic-shelled brachiopods, including both crown and stem groups. However, the complexity, diversity, and biomineralization processes of these microscopic columns are far from clear in brachiopod ancestors. Here, exquisitely well-preserved columnar shell ultrastructures are reported for the first time in the earliest eoobolids Latusobolus xiaoyangbaensis gen. et sp. nov. and Eoobolus acutulus sp. nov. from the Cambrian Series 2 Shuijingtuo Formation of South China. The hierarchical shell architectures, epithelial cell moulds, and the shape and size of cylindrical columns are scrutinised in these new species. Their calcium phosphate-based biomineralized shells are mainly composed of stacked sandwich columnar units. The secretion and construction of the stacked sandwich model of columnar architecture, which played a significant role in the evolution of linguliforms, is highly biologically controlled and organic-matrix mediated. Furthermore, a continuous transformation of anatomic features resulting from the growth of diverse columnar shells is revealed between Eoobolidae, Lingulellotretidae, and Acrotretida, shedding new light on the evolutionary growth and adaptive innovation of biomineralized columnar architecture among early phosphatic-shelled brachiopods during the Cambrian explosion.
Journal Article
Hyolithid-like hyoliths without helens from the early Cambrian of South China, and their implications for the evolution of hyoliths
by
Skovsted, Christian B.
,
Topper, Timothy P.
,
Zhang, Zhifei
in
Analysis
,
Animal Systematics/Taxonomy/Biogeography
,
Biomedical and Life Sciences
2022
Background
A small hyolith, with a triangular operculum and a conical-pyramidal conch with a sharp apex, originally documented as
Ambrolinevitus ventricosus
, is revised based on new material from the Chengjiang biota. The operculum of ‘
Ambrolinevitus
’
ventricosus
displays strong morphological similarities with the operculum of
Paramicrocornus
from the Shuijingtuo Formation (Cambrian Series 2), indicating that the species should be reassigned to
Paramicrocornus
.
Results
Based on the unusual morphology of
Paramicrocornus
, we herein propose a new family Paramicrocornidae fam. nov. A cladistic analysis of Cambrian and Ordovician hyoliths clearly delineates hyolithids as a monophyletic group which evolved from the paraphyletic orthothecids in the early Cambrian and with Paramicrocornidae as its closest relative.
Conclusions
The phylogenetic analysis, together with the distribution of hyoliths from the Cambrian to the Ordovician, reveals the presumptive evolution model of both the skeleton and soft-part anatomy of hyoliths. The Family Paramicrocornidae plays an intermediate role in hyolith evolution, representing the transitional stage in the evolution from orthothecids to hyolithids.
Journal Article
Branched Setae or Attached Macroalgae: A Case Study of an Exceptionally Preserved Brachiopod from the Early Cambrian Chengjiang Lagerstätte
2025
The remarkable conservation of soft tissues within Cambrian fossils has significantly enhanced our comprehension of the origins and evolutionary trajectories of animals, in addition to the progression of ecological intricacy. Here, we report an exceptionally preserved specimen of the lingulid brachiopod Xianshanella haikouensis from the lower Cambrian Chengjiang Lagerstätte, exhibiting branched fringes along the distal ends of its marginal setae. These structures may represent either branched setae or attached macroalgae. The diameter of the branched fringes is slightly larger than that of the marginal setae, and they exhibit third- or fourth-order bifurcations, forming a complex structure comparable in length to the shell. Both the branched fringes and marginal setae are preserved as iron oxides, as revealed by SEM and Micro-XRF analyses, a characteristic preservation mode in Chengjiang fossils. The results of Micro-CT scanning suggest that these branched fringes are preserved along almost the entire distal end of marginal setae. Comparable branched fringes are reminiscent of those found in annelids, and such structural analogs between annelid and brachiopod setae support the homology of brachiopod and annelid setae, representing a lophotrochozoan synapomorphy. An alternative explanation involving attached macroalgae is proposed, given that branched setae have never been documented in either extinct or extant brachiopod taxa. If these structures represent macroalgae, this association could represent a mimicry strategy to deceive predators, although comparable macroalgal fossils remain undocumented in the Chengjiang Lagerstätte. Our research highlights the potential for brachiopod setae to serve roles in sensory function or ecological interactions, offering a new perspective on early animal adaptation and community dynamics.
Journal Article
Combining soft-bodied and three-dimensional fossils to reveal evolutionary modifications in early lingulellotretid brachiopods
by
Topper, Timothy P.
,
Li, Guoxiang
,
Holmer, Lars E.
in
631/181/2468
,
631/181/414
,
Animal Shells - anatomy & histology
2025
Living lingulide brachiopods are traditionally recognised as representatives of evolutionary conservatism, showing little change in general-morphology from their Cambrian ancestors. However, less attention has been given to their anatomical and ontogenetic modifications since their initial appearance. Among these, lingulellotretids are unique, characterized by their typical elongate pedicle foramen and large pseudointerarea. This study describes exquisitely preserved soft-tissue and phosphatic shells of
Lingulellotreta
from Cambrian Series 2 deposits in China and Kazakhstan. Biomineralized novelties in
Lingulellotreta
, including elongate pseudointerarea forming a pouch-like visceral cavity and columnar shell architecture, probably were evolutionarily modified from the unmineralized tubular ancestor
Yuganotheca
during the Cambrian Explosion. Lingulellotretids, however, faced extinction in the Early Ordovician, exemplifying a short-lived evolutionary experiment with a tubular body form in early brachiopods. Since the early Cambrian, lingulide brachiopods have exhibited a long-term evolutionary trend marked by the reduction of pseudointerarea, reflecting a convergence toward a more efficient body plan that ultimately became dominant in later lineages. The intensification of skeletal defences and the increasing demands of filter feeding within benthic communities likely drove these evolutionary modifications and ecological adjustments, culminating in the development of the distinctive, persistent tongue-shaped body of linguloid brachiopods during the Great Ordovician Biodiversification Event.
Tubular lingulellotretid brachiopods, exemplifying a short-lived evolutionary experiment during the Cambrian explosion, gave way to the more efficient tongue-shaped body plan of lingulides by the evolutionary reduction of pseudointerarea.
Journal Article
A fresh look at the Hyolithid Doliutheca from the Early Cambrian (Stage 4) Shipai Formation of the Three Gorges Area, Hubei, South China
2022
New hyolith specimens from the early Cambrian (Stage 4) of the Three Gorges area, western Hubei Province are described and assigned to the species Doliutheca orientalis. Doliutheca are preserved in two taphonomic modes: casts in silty mudstone revealing gross morphology and some soft parts, and internal molds in calcareous pelites, which exhibit new morphological details of the conch and operculum. SEM and Micro-CT analyses show that Doliutheca preserve well-developed platy clavicles and cardinal processes on the interior of the operculum composed of rod-shaped tubular elements. This observation and the distinct cardinal and conical shields of the operculum indicate that Doliutheca could be placed within the Family Paramicrocornidae, most recently established as a group of hyoliths closely related to hyolithids.
Journal Article
Mobergellans from the early Cambrian of Greenland and Labrador: new morphological details and implications for the functional morphology of mobergellans
2018
New morphological features of the mobergellan Discinella micans (Billings, 1871) from the lower Cambrian (Stage 4) of Northeast Greenland and southern Labrador are described. The new features include: (1) the morphology of the larval shell, which is shown to be cap-shaped, subcircular, and with impressions of the internal muscle attachment scars; (2) a range of unusual shell deformations (changes in growth direction resulting in thickened shells, partial detachment of shell laminae and subsequent regrowth, internal projections of shell material increasing the depth of the shell by up to 150%, disturbances and irregular fusion of muscle scars). In addition, we provide new details about the variability in number and shape of the anteriormost internal muscle scars, which often fuse and may vary in number from one to three (resulting in nine to 11 scars in total). Together the new observations provide additional strength for the hypothesis that mobergellan shells represent opercula of an as yet unknown tubular organism.
Journal Article