Shake it off: exploring drivers and outcomes of autotomy in marine invertebrates.

Autotomy refers to self-amputation where the loss of a limb or organ is generally said to be (1) in response to stressful external stimuli; (2) voluntary and nervously mediated; (3) supported by adaptive features that increase efficiency and simultaneously mediate the cost; and (4) morphologically delineated by a predictable breakage plane. It is estimated that this phenomenon has evolved independently nine different times across the animal kingdom, appearing in many different taxa, including vertebrate and invertebrate as well as aquatic and terrestrial animals. Marine invertebrates use this behaviour in a diversity of manners that have yet to be globally reviewed and critically examined. Here, published data from marine invertebrate taxa were used to explore instances of injury as an evolutionary driver of autotomy. Findings suggest that phyla (e.g. Echinodermata and Arthropoda) possibly experiencing high rates of injury (tissue damage or loss) are more likely to be able to perform autotomy. Additionally, this review looks at various morphological, physiological and environmental conditions that have either driven the evolution or maintained the behaviour of autotomy in marine invertebrates. Finally, the use of autotomic abilities in the development of more sustainable and less ecologically invasive fisheries is explored.

HOST SWITCHING IN DICYEMIDS (PHYLUM DICYEMIDA).

Dicyemids (phylum Dicyemida) are the most common and most characteristic endosymbionts in the renal sacs of benthic cephalopod molluscs: octopuses and cuttlefishes. Typically, 2 or 3 dicyemid species are found in a single specimen of the host, and most dicyemids have high host specificity. Host-specific parasites are restricted to a limited range of host species by ecological barriers that impede dispersal and successful establishment; therefore, phylogenies of interacting groups are often congruent due to repeated co-speciation. Most frequently, however, host and parasite phylogenies are not congruent, which can be explained by processes such as host switching and other macro-evolutionary events. Here, the history of dicyemids and their host cephalopod associations were studied by comparing their phylogenies. Dicyemid species were collected from 8 decapodiform species and 12 octopodiform species in Japanese waters. Using whole mitochondrial cytochrome c oxidase subunit 1 (COI) sequences, a phylogeny of 37 dicyemid species, including 4 genera representing the family Dicyemidae, was reconstructed. Phylogenetic trees derived from analyses of COI genes consistently suggested that dicyemid species should be separated into 3 major clades and that the most common genera, Dicyema and Dicyemennea, are not monophyletic. Thus, morphological classification does not reflect the phylogenetic relationships of these 2 genera. Divergence (speciation) of dicyemid species seems to have occurred within a single host species. Possible host-switching events may have occurred between the Octopodiformes and Decapodiformes or within the Octopodiformes or the Decapodiformes. Therefore, the mechanism of dicyemid speciation may be a mixture of host switching and intra-host speciation. This is the first study in which the process of dicyemid diversification involving cephalopod hosts has been evaluated with a large number of dicyemid species and genera.

Structure of the oral tentacles of early ontogeny stage in brachiopod Hemithiris psittacea (Rhynchonelliformea, Rhynchonellida).

Brachiopods have the most complex lophophore in comparison with other lophophorates, i.e., phoronids and bryozoans. However, at early ontogenetic stages, brachiopods have a lophophore of simple morphology, which consists of the oral tentacles. Data on the ultrastructure of the oral tentacles is mostly missing. Nonetheless, it has recently been suggested that the structure of oral tentacles is ancestral for all lophophorates in general, and for brachiopods in particular. The fine structure of the oral tentacles in the brachiopod Hemithiris psittacea is studied using light microscopy, transmission and scanning electron microscopy, cytochemistry and confocal laser scanning microscopy. The oral tentacles have a round shape in transverse section, and four ciliary zones, i.e., one frontal, two lateral, and one abfrontal. Latero-frontal sensory cells occur among the frontal epithelium. Four basiepithelial nerves in the ciliary epithelium are colocalized with ciliary zones. Lophophores of simple morphology in phoronids and brachiopods are characterized by non-specified round forms of tentacles. In phoronids and bryozoans, tentacles have additional latero-frontal ciliary zones that function as a sieve during filtration. In most brachiopods, lateral cilia are involved in the capture of food particles, whereas latero-frontal cells are retained in the frontal zone as sensory elements. The oral tentacles of H. psittacea contain a coelomic canal and have distinct frontal and abfrontal longitudinal muscles, which are separated from each other by peritoneal cells. A similar structure of tentacle muscles occurs in all bryozoans, whereas in phoronids, the frontal and abfrontal tentacle muscles are not separated by peritoneal cells. We suggest that the lophophorates' ancestor had tentacles, which were similar to the tentacles of some phoronids with lophophore of simple morphology. We also assume that the structure of the oral tentacles is ancestral for all brachiopods and the specialization of brachiopod tentacles correlates with the appearance of the double row of tentacles.

Características del hábitat que regulan la estructura de las comunidades de macroinvertebrados en ríos tropicales de montaña (Antioquía, Colombia) / Habitat characteristics that regulate the structure of macroinvertebrate communities in tropical mountain rivers (Antioquía, Colombia)

Introducción: Las comunidades de macroinvertebrados son afectadas simultáneamente por la calidad del agua y las características físicas del hábitat acuático, complicando su uso en la bioindicación. Objetivo: Determinar cuáles variables del hábitat condicionan la comunidad de macroinvertebrados acuáticos en algunas corrientes (quebradas) de montaña del Oriente antioqueño (Colombia). Métodos: El muestreo se realizó en febrero 2021 (periodo de transición seco-lluvia), para evaluar variables físicas y químicas en tres tipos de mesohábitats: rápidos, rizos y pozas en corrientes con coberturas vegetales contrastantes. Los macroinvertebrados fueron recolectados en diez sitios de muestreo con red tipo net, pantalla y manual, y preservados en etanol al 70 %. Resultados: Se recolectaron 4 484 macroinvertebrados (16 órdenes, 46 familias y 75 géneros). El mesohábitat rizo presentó mayores valores de diversidad y abundancia, mientras las pozas presentaron los menores. Hubo diferencias en la concentración de oxígeno, profundidad, velocidad y abundancia de macroinvertebrados entre mesohábitats. Las pozas defirieron de los otros mesohábitats en profundidad, velocidad, así como en la composición, abundancia y riqueza de macroinvertebrados, y fue el hábitat de menor preferencia. Conclusión: La velocidad, profundidad y concentración de oxígeno disuelto, desempeñan un papel muy importante en el establecimiento de las comunidades de macroinvertebrados en los diferentes mesohábitats. En el mismo tipo de mesohábitat, la calidad de la cobertura vegetal determinó la diversidad y abundancia de esta comunidad.

Introduction: Macroinvertebrate communities are affected by water quality and physical characteristics of the aquatic habitat, simultaneously, complicating their use as bioindicators. Objective: To determine which habitat variables regulate the macroinvertebrate community in mountain streams in Eastern of Antioquia (Colombia). Methods: Sampling was carried out in February 2021 (dry-rain transition period), to evaluate physical and chemical variables in three types of mesohabitat: ripples, pools, and rapids in streams with contrasting vegetation covers. The macroinvertebrates were collected from ten sampling sites with a net, screen and manual type net preserved with 70 % ethanol. Results: 4 484 macroinvertebrates were collected (16 orders, 46 families and 75 genera). The ripples mesohabitat presented higher values of diversity and abundance, while the pools presented the lowest. There were differences for oxygen concentration, depth, speed, and macroinvertebrate abundance between mesohabitats. Pools differed from the other mesohabitats in depth, speed, as well as in composition, abundance, and richness in macroinvertebrates, and was the least preferred mesohabitat. Conclusion: Speed, depth, dissolved oxygen concentration played a very important role in the establishment of macroinvertebrates community in different mesohabitats. For the same type of mesohabitat, the quality of the plant cover determined both diversity and abundance of this community.

Composición de los macroinvertebrados según la calidad del agua en turberas de altura, Costa Rica / Composition of macroinvertebrates according to water quality in high altitude peatbogs, Costa Rica

Resumen Introducción: En Costa Rica, las turberas de altura se localizan en la Cordillera de Talamanca de 2 400 a 3 200 m. Se forman a partir de depósitos de materia orgánica en descomposición, con oxígeno reducido y baja disponibilidad de nutrientes. Objetivo: Evaluar la variación en la composición de macroinvertebrados acuáticos según parámetros fisicoquímicos de calidad del agua superficial en turberas de altura, Costa Rica. Métodos: Se colocaron 32 sustratos artificiales en las turberas para ser colonizados por macroinvertebrados. Se realizaron evaluaciones fisicoquímicas mensuales del agua superficial en ocho turberas con visitas mensuales durante la estación lluviosa. Se utilizaron los índices ICA-NSF y BMWP-CR para evaluar la calidad del agua. Resultados: Se registraron un total de diez familias correspondientes a siete géneros. Las familias más representativas fueron Chironomidae y Limnephilidae. La calidad del agua se clasificó según el ICA-NSF como de "calidad media" y como de "calidad regular a mala" según el BMWP-CR. Conclusiones: Las turberas de altura son ecosistemas poco estudiados, esta investigación evidencia la necesidad de diseñar metodologías e índices que evalúen la fauna acuática. Las turberas son diferentes entre sí, y las familias de macroinvertebrados encontrados son tolerantes a las características fisicoquímicas del agua. Es necesario realizar estudios periódicos que evalúen la calidad del agua y su relación con los macroinvertebrados acuáticos para entender la dinámica de las turberas, generar conocimiento e incrementar su protección.

Abstract Introduction: In Costa Rica, high peatbogs are located in the Cordillera de Talamanca from 2 400 to 3 200 m. They are formed from deposits of decomposing organic matter, with reduced oxygen and low nutrient availability. Objective: To evaluate the variation in the composition of aquatic macroinvertebrates according to physicochemical parameters of superficial water quality in high altitude peatbogs, Costa Rica. Methods: 32 artificial substrates were placed in the peatbogs to be colonized by macroinvertebrates. Monthly physicochemical evaluations of surface water were performed in eight peatbogs with monthly visits during the rainy season. The ICA-NSF and BMWP-CR indexes were used to evaluate the water quality. Results: A total of ten families corresponding to seven genera were recorded. The most representative family was Chironomidae and Limnephilidae. Water quality was classified according to the ICA-NSF as "medium quality" and as "regular to poor quality" according to BMWP-CR. Conclusions: The high peatbogs are poorly studied ecosystems; this research shows the need to design methodologies and indices to evaluate the aquatic fauna. The peatbogs are different from each other, and the families of macroinvertebrates found are tolerant to the physicochemical characteristics of the water. Periodic studies that evaluate water quality and its relationship with aquatic macroinvertebrates are necessary to understand the dynamics of peatlands, generate knowledge and increase their protection.

Automatic identification and morphological comparison of bivalve and brachiopod fossils based on deep learning.

Fossil identification is an essential and fundamental task for conducting palaeontological research. Because the manual identification of fossils requires extensive experience and is time-consuming, automatic identification methods are proposed. However, these studies are limited to a few or dozens of species, which is hardly adequate for the needs of research. This study enabled the automatic identification of hundreds of species based on a newly established fossil dataset. An available "bivalve and brachiopod fossil image dataset" (BBFID, containing >16,000 "image-label" data pairs, taxonomic determination completed) was created. The bivalves and brachiopods contained in BBFID are closely related in morphology, ecology and evolution that have long attracted the interest of researchers. We achieved >80% identification accuracy at 22 genera and ∼64% accuracy at 343 species using EfficientNetV2s architecture. The intermediate output of the model was extracted and downscaled to obtain the morphological feature space of fossils using t-distributed stochastic neighbor embedding (t-SNE). We found a distinctive boundary between the morphological feature points of bivalves and brachiopods in fossil morphological feature distribution maps. This study provides a possible method for studying the morphological evolution of fossil clades using computer vision in the future.

ESCAPE PROCESSES IN EMBRYOS OF DICYEMIDS (PHYLUM DICYEMIDA).

Dicyemid mesozoans usually consist of 10 to 40 cells. They are characterized by 2 distinct embryos, vermiform and infusoriform, that develop within the axial cell of the adult. The means of escape of each embryo from the parent body was studied in Dicyema japonicum and Dicyema misakiense, parasites of Octopus sinensis. There were no differences in means of escape between species or embryo type, apparently due to morphological constraints whereby the parents (nematogen or rhombogen) share a similar body organization. Escapes were effected through the gap between adjacent peripheral cells of the adult, rupturing the axial cell membrane and the membrane that envelopes the embryo. After the embryo escaped, the path was closed by the enveloping membrane left behind by the embryo. Vermiform embryos can escape from any region of the body, although more embryos were observed to escape from anterior regions than from posterior regions. Infusoriform embryos escaped from both anterior and posterior regions in the axial cell, with more embryos observed to escape from the posterior regions. The different escape regions for the 2 types of embryo are presumably related to the adult body plan lacking a genital opening, so each different type of embryo has its appropriate site of escape.

A virtuous cycle between invertebrate and robotics research: perspective on a decade of Living Machines research.

Many invertebrates are ideal model systems on which to base robot design principles due to their success in solving seemingly complex tasks across domains while possessing smaller nervous systems than vertebrates. Three areas are particularly relevant for robot designers: Research on flying and crawling invertebrates has inspired new materials and geometries from which robot bodies (their morphologies) can be constructed, enabling a new generation of softer, smaller, and lighter robots. Research on walking insects has informed the design of new systems for controlling robot bodies (their motion control) and adapting their motion to their environment without costly computational methods. And research combining wet and computational neuroscience with robotic validation methods has revealed the structure and function of core circuits in the insect brain responsible for the navigation and swarming capabilities (their mental faculties) displayed by foraging insects. The last decade has seen significant progress in the application of principles extracted from invertebrates, as well as the application of biomimetic robots to model and better understand how animals function. This Perspectives paper on the past 10 years of the Living Machines conference outlines some of the most exciting recent advances in each of these fields before outlining lessons gleaned and the outlook for the next decade of invertebrate robotic research.

Microscopic anatomy of the integument in the ribbon worm Micrura bella (Stimpson, 1857) (Pilidiophora, Nemertea).

The integument of ribbon worms in the order Heteronemertea is distinct from the integuments in the other taxa of nemerteans due to the presence of a special subepidermal glandular layer, the cutis. Among heteronemerteans, the ultrastructure of the cutis has been studied only in the Lineus ruber species complex. In the current study, ultrastructural (transmission electron microscopy) and histochemical studies of the epidermis and the cutis of Micrura bella from the basal Lineage A of the family Lineidae were performed. The epidermis consisted of ciliated and serous gland cells and is separated from the cutis by a layer of the subepidermal extracellular matrix; the basal lamina was not detected. The cutis comprised musculature, two types of mucous and four types of granular gland cells, and pigment cells with four types of granules. In the cutis of juvenile worms, type II granular gland cells and type II mucous cells were not observed. The integument of the caudal cirrus consisted of ciliated and serous gland cells and two intraepidermal lateral nerve cords; the cutis was absent. The compositions of the integument glands of M. bella and the L. ruber species complex are similar, except for the presence of type IV granular gland cells with narrow rod-shaped and lamellated granules exhibiting an alternating dark and light transverse layers and type II mucous cells found only in M. bella.

Altitudinal distribution of the functional feeding groups of aquatic macroinvertebrates using an ecological network in Andean streams / Distribución altitudinal de los grupos funcionales de alimentación de macroinvertebrados acuáticos utilizando una red ecológica en arroyos andinos

Abstract Introduction: Analysis of functional feeding groups (FFG) in aquatic macroinvertebrates is important in understanding the structure, function, and dynamics of ecological processes in ecosystems. Modularity refers to the degree of compartmentalization of food webs and varies between -1 and 1. A network with a modularity value close to 1 is resilient to disturbances and can be interpreted as an indicating factor for the stability of communities. Objective: In this study, we analyzed the trophic structure of benthic macroinvertebrates in La Nitrera stream, the San Juan River, and the Cauca River in the Colombian Andes. Methods: The study was supported by ecological networking techniques using Gephi software. We studied nine sites in dry, rainy, and transition seasons in 2017 and 2018, monitoring changes in the altitude gradient. At each of the sites, the organisms were captured and determined, and physicochemical and hydraulic information was obtained. Results: The variance component analysis allowed to explain the variability of the data by relating the following environmental variables: FFG, diversity, richness, modularity, season, and time. Simple multifactorial ANOVA indicated that significant changes in FFG were associated with altitude, and modularity to time. The allocation of the FFG was done by stomach analysis and secondary information. Conclusion: The transition season had the highest modularity, possibly due to the recolonization of some biotopes caused by the decrease in the velocity of water currents. La Nitrera and San Juan presented higher values than the Cauca, which may indicate that the altitudinal change and velocity of water currents affects the compartmentalization of the network.

Resumen Introducción: El análisis de grupos funcionales de alimentación (GFA) en macroinvertebrados acuáticos es importante para comprender la estructura, función y dinámica de los ecosistemas de procesos ecológicos. La modularidad se refiere al grado de compartimentación de las redes alimentarias y varía entre -1 y 1. Una red con un valor de modularidad cercano a 1 es resistente a las alteraciones y puede interpretarse como un factor indicativo para la estabilidad de las comunidades. Objetivo: En este estudio se analizó la estructura trófica de los macroinvertebrados bentónicos, un elemento importante en la calidad ambiental, en el arroyo La Nitrera, el río San Juan y el río Cauca. Métodos: El estudio contó con el apoyo de técnicas de redes ecológicas utilizando el software Gephi. En 2017 y 2018, estudiamos nueve sitios en estaciones secas, lluviosas y de transición, monitoreando cambios en el gradiente de altitud. En cada uno de los sitios se capturaron y determinaron los organismos y se recogió información fisicoquímica e hidráulica. Resultados: El análisis de componentes de varianza permitió explicar la variabilidad de los datos relacionando las siguientes variables ambientales: GFA, diversidad, riqueza, modularidad, estación y tiempo. La ANOVA simple multifactorial indicó que existen cambios significativos en los GFA en relación con la altitud, y la modularidad con el tiempo. La asignación de los GFA se realizó mediante análisis estomacal e información secundaria. Conclusión: La temporada de transición tuvo la mayor modularidad, posiblemente debido a la recolonización de algunos biotopos provocada por la disminución de la velocidad del cauce. La Nitrera y San Juan presentaron valores superiores a los del Cauca, lo que puede indicar que el cambio altitudinal y la velocidad de las corrientes de agua influyen en la compartimentación de la red.

A Cambrian tommotiid preserving soft tissues reveals the metameric ancestry of lophophorates.

Among extant animals, Lophotrochozoa accounts for the majority of phyla.1 This bilaterian clade radiated rapidly during the Cambrian explosion, obfuscating its phylogenetic relationships and rendering many aspects of its early evolution uncertain. Many early lophotrochozoans are known only from isolated skeletal microfossils, "small shelly fossils," often derived from larger animals with complex multi-element skeletons.2 The discovery of articulated fossils has revealed surprising insights into the animals from which these skeletal pieces were derived, such as paired shells in the mollusc Halkieria.3 Tommotiids are a key group of phosphatic early skeletal fossils that first appear in the late early Cambrian.4,5 Although their affinities were previously obscure, discoveries of partial scleritomes and investigations of growth and microstructure6 provide links with Brachiopoda7,8 and Phoronida,9 two of the lophophorate phyla. By contrast, the body plan of camenellan tommotiids remains a palaeontological mystery, with hypothetical reconstructions representing motile, benthic, dorsally armored worms.4,10 Here, we describe an articulated camenellan (Wufengella bengtsoni gen. et sp. nov.) from the Cambrian Chengjiang Biota, China, revealing the morphology of the scleritome and the first soft tissues from an adult tommotiid. Wufengella carries two dorsal rows of sclerites in a highly asymmetric arrangement, flanked by smaller, cap-shaped sclerites. The scleritome was fringed by iterated fascicles of chaetae and two layers of flattened lobes. Phylogenetic analysis confirms that camenellans occupy a deep branch in lophophorate phylogeny, prior to the acquisition of a sessile lifestyle. Wufengella reveals direct evidence for a metameric body plan reminiscent of annelids early in the evolutionary history of lophophorates.11,12.

A new species of Echinoderes (Kinorhyncha: Cyclorhagida: Echinoderidae) from the Argentinean continental shelf with notes on its postembryonic development and on subcuticular morphological characters unreported for Kinorhyncha.

Echinoderes goku sp. nov. is described from adult and juvenile specimens in samples collected from the Argentinean continental shelf in March 2019 during the campaign of the Motorsailer Bernardo Houssay, based on light and scanning electron microscopy studies. Echinoderes goku sp. nov. is differentiated from other species by its unique spine and tube pattern: (1) middorsal spine on segments 4, 6, and 8, (2) spine in lateroventral position on segments 69, (3) tube present in subdorsal position on segment 2, sublateral to midlateral position on segments 7 and 9, lateral accessory position on segment 8, lateroventral position on segment 5 and ventrolateral position on segment 2. Additionally, this species (4) lacks type-2 gland cell outlets and female-specific papillae or subcuticular funnel-like structures. (5) The male possesses a sublateral tube on segment 10 with a basal cylindrical reinforcement structure and three penile spines on segment 11, whereas the female lacks penile spines but reveals a lateral terminal accessory spine on segment 11 and on segment 10 lacks a subdorsal tube but possesses instead a sublateral fringe on the surface connected with a subcuticular conical structure and a basal cylindrical reinforcement structure. Among several juvenile specimens, one was captured in the process of moulting from the last juvenile stage to the female. Whereas the last juvenile stage revealed a sublateral tube on segment 10, the moulting female lacked this tube and showed a conical subcuticular structure with a basal cylindrical reinforcement structure instead. A cylindrical reinforcement structure at the base of a tube and a subcuticular conical structure have not been reported before for Kinorhyncha, and 14 selected species of Echinoderidae available for study were screened for these characters in order to get a first idea of their possible taxonomic importance. This article reports about the second species of Echinoderes from Argentinean waters.

Tentacle muscles in brachiopods: Ultrastructure and relation to peculiarities of life style.

Although the morphology of the brachiopod tentacle organ, the lophophore, is diverse, the organization of tentacles has traditionally been thought to be similar among brachiopods. We report here, however, that the structure of the tentacle muscles differs among brachiopod species representing three subphyla: Lingula anatina (Linguliformea: Linguloidea), Pelagodiscus atlanticus (Linguliformea: Discinoidea), Novocrania anomala (Craniiformea), and Coptothyris grayi (Rhynchonelliformea). Although the tentacle muscles in all four species are formed by myoepithelial cells with thick myofilaments of different diameters, three types of tentacle organization were detected. The tentacles of the first type occur in P. atlanticus, C. grayi, and in all rhynchonelliforms studied before. These tentacles have a well-developed frontal muscle and a small abfrontal muscle, which may reflect the ancestral organization of tentacles of all brachiopods. This type of tentacle has presumably been modified in other brachiopods due to changes in life style. Tentacles of the second type occur in the burrowing species L. anatina and are characterized by the presence of equally developed smooth frontal and abfrontal muscles. Tentacles of the third type occur in N. anomala and are characterized by the presence of only well-developed frontal muscles; the abfrontal muscles are reduced due to the specific position of tentacles during filtration and to the presence of numerous peritoneal neurites on the abfrontal side of the tentacles. Tentacles of the first type are also present in phoronids and bryozoans, and may be ancestral for all lophophorates.

Intrabody Tetrodotoxin Distribution and Possible Hypothesis for Its Migration in Ribbon Worms Cephalothrix cf. simula (Palaeonemertea, Nemertea).

Tetrodotoxin (TTX) is a potent neurotoxin found in many marine and terrestrial animals, but only a few species, such as the ribbon worms of the genus Cephalothrix, accumulate it in extremely high concentrations. The intrabody distribution of TTX in highly toxic organisms is of great interest because it helps researchers to understand the pathways by which the toxin migrates, accumulates, and functions in tissues. Using immunohistochemistry with anti-TTX antibodies, the authors of this study investigated the toxin's distribution inside the organs, tissues, and cells of Cephalothrix cf. simula. The cell types of TTX-positive tissues were identified by light microscopy. The main sites of TTX accumulation occurred in the secretory cells of the integuments, the microvilli of the epidermal ciliary cells, cephalic glands, the glandular epithelia of the proboscises, the enterocytes of the digestive systems, and nephridia. Obtained data suggest the toxin migrates from the digestive system through blood vessels to target organs. TTX is excreted from the body through the nephridia and mucus of epidermal cells.

Environmental DNA gives comparable results to morphology-based indices of macroinvertebrates in a large-scale ecological assessment.

Anthropogenic activities are changing the state of ecosystems worldwide, affecting community composition and often resulting in loss of biodiversity. Rivers are among the most impacted ecosystems. Recording their current state with regular biomonitoring is important to assess the future trajectory of biodiversity. Traditional monitoring methods for ecological assessments are costly and time-intensive. Here, we compared monitoring of macroinvertebrates based on environmental DNA (eDNA) sampling with monitoring based on traditional kick-net sampling to assess biodiversity patterns at 92 river sites covering all major Swiss river catchments. From the kick-net community data, a biotic index (IBCH) based on 145 indicator taxa had been established. The index was matched by the taxonomically annotated eDNA data by using a machine learning approach. Our comparison of diversity patterns only uses the zero-radius Operational Taxonomic Units assigned to the indicator taxa. Overall, we found a strong congruence between both methods for the assessment of the total indicator community composition (gamma diversity). However, when assessing biodiversity at the site level (alpha diversity), the methods were less consistent and gave complementary data on composition. Specifically, environmental DNA retrieved significantly fewer indicator taxa per site than the kick-net approach. Importantly, however, the subsequent ecological classification of rivers based on the detected indicators resulted in similar biotic index scores for the kick-net and the eDNA data that was classified using a random forest approach. The majority of the predictions (72%) from the random forest classification resulted in the same river status categories as the kick-net approach. Thus, environmental DNA validly detected indicator communities and, combined with machine learning, provided reliable classifications of the ecological state of rivers. Overall, while environmental DNA gives complementary data on the macroinvertebrate community composition compared to the kick-net approach, the subsequently calculated indices for the ecological classification of river sites are nevertheless directly comparable and consistent.

Mechanical adaptation of brachiopod shells via hydration-induced structural changes.

The function-optimized properties of biominerals arise from the hierarchical organization of primary building blocks. Alteration of properties in response to environmental stresses generally involves time-intensive processes of resorption and reprecipitation of mineral in the underlying organic scaffold. Here, we report that the load-bearing shells of the brachiopod Discinisca tenuis are an exception to this process. These shells can dynamically modulate their mechanical properties in response to a change in environment, switching from hard and stiff when dry to malleable when hydrated within minutes. Using ptychographic X-ray tomography, electron microscopy and spectroscopy, we describe their hierarchical structure and composition as a function of hydration to understand the structural motifs that generate this adaptability. Key is a complementary set of structural modifications, starting with the swelling of an organic matrix on the micron level via nanocrystal reorganization and ending in an intercalation process on the molecular level in response to hydration.

The nervous system of the most complex lophophore provides new insights into the evolution of Brachiopoda.

The lophophore is a tentacle organ unique to the lophophorates. Recent research has revealed that the organization of the nervous and muscular systems of the lophophore is similar in phoronids, brachiopods, and bryozoans. At the same time, the evolution of the lophophore in certain lophophorates is still being debated. Innervation of the adult lophophore has been studied by immunocytochemistry and confocal laser scanning microscopy for only two brachiopod species belonging to two subphyla: Linguliformea and Rhynchonelliformea. Species from both groups have the spirolophe, which is the most common type of the lophophore among brachiopods. In this study, we used transmission electron microscopy, immunocytochemistry, and confocal laser scanning microscopy to describe the innervation of the most complex lophophore (the plectolophe) of the rhynchonelliform species Coptothyris grayi. The C. grayi lophophore (the plectolophe) is innervated by three brachial nerves: the main, second accessory, and lower. Thus, the plectolophe lacks the accessory brachial nerve, which is typically present in other studied brachiopods. All C. grayi brachial nerves contain two types of perikarya. Because the accessory nerve is absent, the cross nerves, which pass into the connective tissue, have a complex morphology: each nerve consists of two ascending and one descending branches. The outer and inner tentacles are innervated by several groups of neurite bundles: one frontal, two lateral, two abfrontal, and two latero-abfrontal (the latter is present in only the outer tentacles). Tentacle nerves originate from the second accessory and lower brachial nerves. The inner and outer tentacles are also innervated by numerous peritoneal neurites, which exhibit acetylated alpha-tubulin-like immunoreactivity. The nervous system of the lophophore of C. grayi manifests several evolutionary trends. On the one hand, it has undergone simplification, i.e., the absence of the accessory brachial nerve, which is apparently correlated with a reduction in the complexity of the lophophore's musculature. On the other hand, C. grayi has a prominent second accessory nerve, which contains large groups of frontal perikarya, and also has additional nerves extending from the both ganglia to the medial arm; these features are consistent with the complex morphology of the C. grayi plectolophe. In brachiopods, the evolution of the lophophore nervous system apparently involved two main modifications. The first modification was the appearance and further strengthening of the second accessory brachial nerve, which apparently arose because of the formation of a double row of tentacles instead of the single row of the brachiopod ancestor. The second modification was the partial or complete reduction of some brachial nerves, which was correlated with the reduced complexity of the lophophore musculature and the appearance of skeletal structures that support the lophophore.

From the individual to the colony: Marine invertebrates as models to understand levels of biological organization.

The developmental and evolutionary principles of coloniality in marine animals remain largely unexplored. Although many common traits have evolved independently in different groups of colonial animals, questions about their significance for colonial life histories remain unanswered. In 2018 (Nov. 25 - Dec. 8), the inaugural course on the Evolution of Coloniality and Modularity took place at the Center for Marine Biology of the University of São Paulo (CEBIMAR-USP), Brazil. During the intensive two-week graduate-level course, we addressed some of the historical ideas about animal coloniality by focal studies in bryozoans, tunicates, cnidarians, and sponges. We discussed many historical hypotheses and ways to test these using both extant and paleontological data, and we carried direct observations of animal colonies in the different phyla to address questions about coloniality. We covered topics related to multi-level selection theory and studied colonial traits, including modular miniaturization, polymorphism, brooding, and allorecognition. Course participants carried out short research projects using local species of animals to address questions on allorecognition and regeneration in ascidians and sponges, fusion and chimerism in anthoathecate hydrozoans, and evolution of polymorphism in bryozoans. Although many questions remain unanswered, this course served as a foundation to continue to develop a developmental and evolutionary synthesis of clonal and modular development in colonial marine organisms.

Exceptionally preserved early Cambrian bilaterian developmental stages from Mongolia.

Fossilized invertebrate embryonic and later developmental stages are rare and restricted largely to the Ediacaran-Cambrian, providing direct insight into development during the emergence of animal bodyplans. Here we report a new assemblage of eggs, embryos and bilaterian post-embryonic developmental stages from the early Cambrian Salanygol Formation of Dzhabkan Microcontinent of Mongolia. The post-embryonic developmental stages of the bilaterian are preserved with cellular fidelity, possessing a series of bilaterally arranged ridges that compare to co-occurring camenellan sclerites in which the initial growth stages retain the cellular morphology of modified juveniles. In this work we identify these fossils as early post-embryonic developmental stages of camenellans, an early clade of stem-brachiopods, known previously only from isolated sclerites. This interpretation corroborates previous reconstructions of camenellan scleritomes with sclerites arranged in medial and peripheral concentric zones. It further supports the conjecture that molluscs and brachiopods are descended from an ancestral vermiform and slug-like bodyplan.

Anatomy of the coelomic system in Novocrania anomala (Brachiopoda, Craniiformea) and relationships within brachiopods.

Brachiopoda is a relict phylum of marine benthic animals that have not been adequately studied with modern microscopy methods. Microscopic study may provide useful information on the evolution of the brachiopod body plan and brachiopod phylogeny. Understanding the organisation of the coelomic system is important because of its role in body form and compartmentalisation. Most brachiopods are considered to have a bipartite coelomic system; the only known exception is Lingulida, which have a tripartite coelomic system. In the present study, we provide the first complete 3D reconstruction of the coelomic system in the craniide brachiopod Novocrania anomala (Müller, 1776). Its coelomic system consists of the following five main parts, which are entirely separated from each other: 1) a pair of large brachial canals; 2) a complex system of paired small brachial canals and a perioesophageal coelom; 3) frontal coelomic chambers; 4) a main trunk coelom, which includes several semi-detached muscular chambers and mantle sinuses; and 5) a pair of posterior adductors chambers. These results indicate that the coelomic system of N. anomala (and perhaps of other craniides) is complex and cannot be considered to be bipartite or tripartite. The frontmost part of the coelomic system is represented by a pair of frontal chambers, which are considered to be a part of the lophophore but which are derived from dorsal mantle fold extensions and thus may be a part of the trunk coelomic system. A number of similarities were discovered between craniiformean and rhynchonelliformean coelomic systems, including the prominent dorsal projections of the large brachial canals and the morphological features of the perioesophageal coelom. The complex subdivision of the N. anomala trunk coelom is explained by the location and function of muscles, and by the location of several mesenteries.