The Aestivation Chamber of the Giant Earthworm Glossoscolex bergi (Glossoscolecidae) in the Subtropical Rainforest of Misiones (Argentina)

Glossoscolex bergi (Glossoscolecidae) is a giant earthworm from the rainforest of Misiones (Argentina). The large size of its aestivation chamber and the meniscate burrows connected to it allow us to describe morphological details and reinterpret some characteristics of the fossil counterpart Castrichnus incolumis. The concavity of menisci, either in the burrow or in those pellets lining the chamber, shows concentric ridges and radiating striae that result from the impression of the last segments of the body. The meniscate burrows associated to the chamber are different from described ichnospecies of Taenidium, because of the surface texture of the meniscus. The aestivation chamber was produced during an atypical four-month drought in a region that lacks a seasonal climate. This suggests that Castrichnus incolumis would be also an indicator of drought periods even in non seasonal climates.     

Is the aestivation of the earthworm Hormogaster elisae a paradiapause?

Abstract. We report the results of a laboratory study of the aestivation of Hormogaster elisae to determine the nature of the inactive period (diapause or quiescence) and to evaluate the influence of soil moisture, temperature, season, and earthworm body weight on the process. The results showed that specimens of H. elisae underwent facultative diapause—paradiapause—characterized by the construction of aestivation chambers in which the animal coils up and its activity decreases. Soil moisture appeared to be the most important environmental factor involved in the onset of aestivation. Temperature and time of year also had some influence, but earthworm body weight appeared to have none. Aestivating earthworms showed a decrease of 41.6% in mean body weight. Once replaced in soil with 20% moisture content, they took 6.4±3.1 d to exit their chambers and another 6.5±3.6 d to recover their initial body weight.     

Teisseirei barattinia Roselli : the first sphinx moth trace fossil from palaeosols,and its distinct type of wall

The trace fossil Teisseirei barattinia, found in Cenozoic formations of Uruguay and Argentina, is an elongated chamber recognizable by its depressed cross‐section, antechamber, and its multi‐layered lining with an inner surface texture composed of densely packed sub‐rectangular to sub‐triangular pits. Our recent behavioural observations on larval and pupal stages of Sphingidae (Lepidoptera), particularly on Manduca rustica, suggest that Teisseirei barattinia is the pupation chamber of a sphinx moth. Last instar larvae of Manduca rustica, Eumorpha anchemolus and E. labruscae were placed in terraria to observe their burrowing behaviour and to recover pupation chambers. Chambers show depressed or plane convex cross‐sections as T. barattinia. The internal surface texture of walls is also similar to that of T. barattinia. The same pattern could be obtained experimentally by pressing the true legs of Manduca rustica larva against plasticine. The multi‐layered wall structure, shown by T. barattinia, is a new type of lining for insect trace fossils in palaeosols that result from soil packing combined with discharges of abundant liquid excretion by soft‐bodied larvae, as in the case of Manduca rustica. T. barattinia is the first trace fossil documented in palaeosols attributed to sphinx moths and supported by macro and micromorphological comparisons with extant pupation chambers. The shallow emplacement of moth pupation chambers in soils suggests that T. barattinia would be a good indicator of palaeosol upper horizons.     

CLEPTOPARASITISM AND DETRITIVORY IN DUNG BEETLE FOSSIL BROOD BALLS FROM PATAGONIA, ARGENTINA

Abstract:  Traces within traces is a new ichnological field that is meant to shed light on significative palaeoecological aspects. Dung beetle fossil brood balls ( Coprinisphaera ispp.), from the Middle Eocene – Lower Miocene Sarmiento Formation of Patagonia, Argentina, show two different trace fossils excavated in its infillings and/or wall that reveal the presence and relationships among different components of past dung communities. Tombownichnus pepei n. isp. is represented by elongated pits, circular to elliptical in cross-section, occurring in the centre or beside ovoid mounds in the internal surface of the Coprinisphaera wall. These traces record the activity of cleptoparasites, such as other dung beetles or flies, whose larvae were probably carried passively with the dung for provisions. Tombownichnus pepei would represent the pupation chambers excavated by full grown larvae in the Coprinisphaera wall after completing their development inside provisioned dung. The other trace fossil, Lazaichnus fistulosus is represented by circular to subcircular borings occurring in Coprinisphaera walls, in connection with an internal gallery in their infillings. Its connection also with meniscate burrows and chambers in the surrounding palaeosol attributable to aestivation chambers of earthworms revealed that these organisms would have been active cleptoparasites or detritivores in dung beetle fossil brood balls.     

Minerals in the rhizosphere: overlooked mediators of soil nitrogen availability to plants and microbes

Non-native earthworms are a continued source of environmental change in the northeastern United States that may affect trace metals in the plant-soil system, with largely unknown effects. We assessed earthworm impacts on exchangeable and strong acid extractable (total) concentrations and pools of Al, Fe, Cu, Zn, Mo, Pb in non-point source polluted, forest soil horizons (Organic, A, and B) and foliar metals concentrations in young (<?3 years) Acer saccharum and Polystichum acrostichoides at four proximal forests in the Finger Lakes Region of New York. We observed decreasing total trace metal Organic horizon pools and increasing total trace metal A horizon concentrations as a function of increasing earthworm biomass. Earthworms had limited effects on exchangeable concentrations in A and B horizons and total metal concentrations in the B horizon. Foliar trace metal concentrations in Acer were better explained by earthworm biomass than soil concentrations but foliar concentrations for Polystichum were poorly predicted by both earthworm biomass and soil metal concentrations. Our results suggest that earthworms can affect trace metal uptake by some plants, but not by increasing soil trace metal exchangeability or from changing soil properties (pH, %SOM, or cation exchange capacity). Instead, non-native earthworms may indirectly alter understory plant uptake of trace metals.     

Rhabditis terricola: An opportunistic nematode parasite of earthworm cocoons

During a study on the breeding rate of the earthworms Lumbricus rubellus and Eisenia foetida, nematodes identified as Rhabditis terricola were found consistently in large numbers in earthworm cocoons that failed to hatch after an adequate incubation period. This nematode species, which was previously known as a saprophyte, was found to invade earthworm cocoons and reproduce within, causing extensive productivity losses in earthworm cultures. In this study, R. terricola was effectively eradicated from earthworm cultures by rinsing the earthworms in tap water and transferring them repeatedly to sterile bedding.     

Fossil bee nests,coleopteran pupal chambers and tuffaceous paleosols from the Late Cretaceous Laguna Palacios Formation,Central Patagonia (Argentina)

The Late Cretaceous Laguna Palacios Formation in Central Patagonia (San Jorge Basin), southern South America, is composed of tuffaceous deposits supplied by periodical volcanic ash falls partly reworked by rivers, on broad plains. Variations in ash-fall rates allowed the formation of stacked, mature paleosols, which are one of the most characteristic features of this formation. The mature paleosols show well-developed horizons, ped structure and bear an intricate network of trace fossils mostly produced by small roots and invertebrates. Two different insect trace fossils could be recognized in this formation: sweat bee nests and coleopteran pupal chambers. Fossil bee nests are composed of inclined tunnels with cells attached to them by means of short necks, a typical construction of bees of the subfamily Halictinae. Similar halictine constructions were reported from the Cretaceous of the USA. Coleopteran pupal chambers are discrete, ovoid structures, having an internal cavity with a smooth surface, and an outer wall of lumpy appearance composed of different layers of soil material. They are commonly constructed by the larvae of different families of Coleoptera. Similar trace fossils were previously reported from the Asencio Formation (Late Cretaceous–Early Tertiary) of Uruguay and from the Djadokhta Formation of Mongolia (Late Cretaceous). These trace fossils constitute some of the only paleontological data from the Laguna Palacios Formation, allowing inferences about its paleoecology, paleoclimatology and paleogeography. Ecological preferences of Halictinae, as well as some features of the nests, suggest a temperate, seasonal climate and an environment dominated by low vegetation for the Laguna Palacios Formation, which is also compatible with sedimentologic and pedogenic evidence. The morphology of the nests, typical of North American halictinae, adds more evidence to the hypothesis of the existence of faunal interchange between North and South America by the Late Cretaceous. The fossil nests constitute some of the oldest evidence of bees in the fossil record, the third known record of bees of Cretaceous age and the first for the Southern Hemisphere. The two traces described are, together with those of Dakota and the Gobi Desert, the only trace fossils from paleosols of Cretaceous age that can be certainly attributed to insects.     

White-tailed deer (Odocoileus virginianus) fecal pellet decomposition is accelerated by the invasive earthworm Lumbricus terrestris

Exotic European earthworms have expanded into worm-free forests of the United States. Concurrently, populations of the white-tailed deer, Odocoileus virginianus, have also increased. During winter, deer use hemlock stands for cover while browsing elsewhere, creating a net organic matter flux into these stands. Deer fecal pellets can provide annual inputs of 48.1 kg C, 1.4 kg N, and 1.3 kg Ca per hectare. We tested the hypothesis that these pellets were readily consumed by invading earthworms. The litter-feeding anecic earthworm Lumbricus terrestris redistributed fecal pellets and accelerated mass and nutrient loss rates. These losses are likely due to the combination of enhanced fragmentation and decomposition as earthworms drag pellets into their burrows for consumption. This nutrient subsidy may be an important source of high quality “litter” input to hemlock stands, which may in turn facilitate the invasion of these stands by earthworms under high deer densities.     

Earthworms in the diet of Herring Gulls Larus argentatus breeding on an off-shore island

ABSTRACT

Pellets of indigestible material regurgitated by Herring Gulls Larus argentatus breeding on Lady Isle, Firth of Clyde, Scotland, were collected in 2018 and 2019 and examined for earthworm chaetae. Nearly two-thirds (65.6%) of the 314 pellets came from gulls that had consumed one or more earthworms. Significantly fewer pellets (57.6%) contained chaetae in 2018, a relatively dry May to July, than in 2019 (72.0%) when rainfall was close to the long-term average for May to July. There were significant associations between the presence of large quantities of terrestrial vegetation in the pellet and the detection of large numbers of earthworm chaetae and/or fragments of terrestrial arthropods, suggesting that recent consumption of these food items is likely when pellets contain large quantities of vegetation.     

Sine Systemate Chaos? A Versatile Tool for Earthworm Taxonomy: Non-Destructive Imaging of Freshly Fixed and Museum Specimens Using Micro-Computed Tomography

In spite of the high relevance of lumbricid earthworms (‘Oligochaeta’: Lumbricidae) for soil structure and functioning, the taxonomy of this group of terrestrial invertebrates remains in a quasi-chaotic state. Earthworm taxonomy traditionally relies on the interpretation of external and internal morphological characters, but the acquisition of these data is often hampered by tedious dissections or restricted access to valuable and rare museum specimens. The present state of affairs, in conjunction with the difficulty of establishing primary homologies for multiple morphological features, has led to an almost unrivaled instability in the taxonomy and systematics of certain earthworm groups, including Lumbricidae. As a potential remedy, we apply for the first time a non-destructive imaging technique to lumbricids and explore the future application of this approach to earthworm taxonomy. High-resolution micro-computed tomography (μCT) scanning of freshly fixed and museum specimens was carried out using two cosmopolitan species, Aporrectodea caliginosa and A. trapezoides. By combining two-dimensional and three-dimensional dataset visualization techniques, we demonstrate that the morphological features commonly used in earthworm taxonomy can now be analyzed without the need for dissection, whether freshly fixed or museum specimens collected more than 60 years ago are studied. Our analyses show that μCT in combination with soft tissue staining can be successfully applied to lumbricid earthworms. An extension of the approach to other families is poised to strengthen earthworm taxonomy by providing a versatile tool to resolve the taxonomic chaos currently present in this ecologically important, but taxonomically neglected group of terrestrial invertebrates.     

Scolicia and its producer in shallow-marine deposits of the Miocene Chenque Formation (Patagonia,Argentina): functional morphology and implications for understanding burrowing behavior

Abstract

Scolicia is one of the most conspicuous trace fossils in lower shoreface deposits of the Miocene Chenque Formation of Patagonia, Argentina. This ichnotaxon consists of horizontal, sinuous or meandering trace fossils with a laminated backfill and two parallel strings located at the base. Abundant body fossils attributed to Brisaster iheringi occur in close association to these trace fossils. The echinoids are very well preserved, and most of the specimens have some areas with their spines attached in life position. In particular, preservation of the subanal tufts of spines, associated to isopores (which are interpreted as being associated to funnel building tube feet), supports the interpretation that these organisms are the producers of Scolicia isp. A shaft connecting the burrow with the sediment-water interface was not observed in vertical sections of the excavations, although on bedding plane surfaces, some small circular tube outlets are recognized. However, the great diversity and abundance of ichnofossils co-occurring in these deposits may preclude adequate identification of vertical structures. These deposits provide an excellent opportunity to integrate trace fossil data, body fossil information and observations from modern analogues in order to perform a morpho-functional analysis of Scolicia.     

Determination of the diet of the stone curlew (Bwhinus oedicnemus) by faecal analysis

The value of faecal analysis for estimating the composition of the diet of stone curlews was assessed by examination of the faeces of a captive bird fed on a measured diet. Remains of soft–bodied prey were under–represented in the faeces but the results could be adjusted to allow for this. Estimation of the size of prey was possible by measurement of remnants such as earthworm chaetae and insect mandibles. In some types of prey large specimens left more remnants than small ones and a correction was devised to allow for this. Regurgitated pellets contained large, hard remains and there was no trace of some important prey types such as earthworms. A method is described for estimating the composition of the diet of the wild stone curlews from counts of prey remains in their faeces.     

Interactions between Nematodes and Earthworms: Enhanced Dispersal of Steinernema carpocapsae

Dispersal of the nematode Steinernema carpocapsae (All strain), applied on the top or the bottom of soil columns, was tested in the presence or absence of two earthworm species, Lumbricus terrestris or Aporrectodea trapezoides. Nematode dispersal was estimated after a 2-week period with a bioassay against the greater wax moth, Galleria mellonella. Vertical dispersal of nematodes was increased in the presence of earthworms. When nematodes were placed on the surface of soil columns, significantly more nematodes dispersed to the lower half of the columns when either earthworm species was present than when earthworms were not present. When nematodes were placed on the bottom of soil columns, significantly more nematodes dispersed to the upper half of the columns when L. terrestris was present than when A. trapezoides was present or in the absence of earthworms. Because nematodes were found on the exterior and in the interior of earthworms, nematode dispersal may be enhanced by direct contact with the earthworms.     

Local distribution of native and invasive earthworms and effects on a native salamander

North America is home to both native and invasive earthworms acting as ecosystem engineers as they build burrows that can serve as habitat for other species or otherwise alter soil structure, affecting nutrient cycling and other ecosystem processes. Here I determine where and what earthworm species commonly occur in my study area, and compare effects of native and invasive earthworms on the common woodland salamander, Plethodon cinereus, in field surveys and laboratory experiments. The native earthworm Eisenoides carolinensis was the most common earthworm, followed by two invasive species Dendrobaena octaedra and Octolasion tyrtaeum. The presence of O. tyrtaeum was associated with a narrower O-horizon (i.e., organic layer in the soil). Using structural equation modeling to explore direct and indirect pathways of these three most common earthworm species on salamanders, I found O. tyrtaeum occurrence was negatively correlated with nighttime salamander counts, a proxy for total salamander numbers, mediated by negative effects on O-horizon depth and microinvertebrate numbers. In the laboratory, O. tyrtaeum and D. octaedra consumed more leaf litter per gram of earthworm per day than the native E. carolinensis. However, salamanders consumed earthworms and used burrows of all native and invasive species of earthworms similarly. The potential for negative indirect effects of the invasive earthworm O. tyrtaeum on P. cinereus was demonstrated both in the field and laboratory, highlighting that seemingly small differences between native and invasive ecosystem engineers have the potential to significantly alter the effects of these closely related native and invasive organisms.     

A comparative study of gut microbiota profiles of earthworms fed in three different substrates

The gut microbiome of earthworms has a complex interdependence with the host. When the soil minerals pass through earthworm’s gut, they may affect the gut microbiota. To gain insight into the response of gut microbiota to the passed minerals, we fed earthworm (Eisenia fetida) on nutrient-poor soil and ore powder, and used high throughput sequencing to characterize the earthworm intestinal microbial community to find evidence for a core bacterial community of the E. fetida. The results showed that earthworms’ gut maintained a core microbiome that appeared in all samples. These core microbiota may play a significant role in a species’ environmental interactions. The composition of intestinal microbiomes varied with substrates. The earthworm guts from two nutrient-poor substrates had similar microbial communities and they were different from nutrient-rich substrate. Proteobacteria and Bacteroidetes were more abundant in the gut of earthworms kept on a nutrient-poor substrate such as ore powder or mineral soil than in the gut of earthworms kept in organic-rich compost soil; some of these microorganisms may help earthworms survive in nutrient-poor substrates.     

Earthworm communities a bioindicator of arable land management practices: A case study in semiarid region of India

The abundances and activities of earthworms in arable lands depend strongly on management practices and; therefore earthworm can act as a potential bioindicator of land use practices. In this study earthworm diversity and abundances was measured in three differently managed agro-ecosystems i.e. conventional, integrated and organic, in some sites of northern semiarid region of India. The maximal numbers of earthworm occurrence were in integrated farming system (100%, all studied sites showed the presence) followed by organically managed (70%) and conventional (18.9%) agro-ecosystems. A total of six species belonging to four different families were identified during this survey and all recorded earthworm species were present in organically managed agro-ecosystems, while integrated and conventional arable lands exhibited only three earthworm species. The abundance of earthworms in arable system was also directly related to the management practices. The values of ecological indices e.g. Shannon diversity (H′), species dominance (C), the species richness (S) and evenness (E) indicated the anthropogenic pressure on earthworm communities in arable lands of northern India.     

Local-scale correlates of native and non-native earthworm distributions in juniper-encroached tallgrass prairie

European and Asian earthworms have invaded much of North America with profound impacts to soils, plant communities, and animal populations. However, few studies have assessed local-scale correlates of earthworm distributions, and most invasive earthworm research has occurred in northern forests. Additionally, despite several studies showing facilitative relationships between invasive earthworms and invasive plants, no research has assessed a potential facilitative interaction between earthworms and woody plants encroaching into prairies. We conducted the first assessment of factors influencing local-scale distributions of native and non-native earthworms for the U.S. Great Plains in a tallgrass prairie-woodland mosaic experiencing eastern redcedar (Juniperus virginiana) encroachment. We documented both native and non-native earthworms, including non-native species from Eurasia (Aporrectodea spp.) and South America (Family Ocnerodrilidae). Native and non-native earthworm distributions were strongly correlated, yet local-scale predictors of distribution also differed between the groups. Native earthworms were more likely to occur near roads and in areas with moist soils. Contrary to expectation, we found no evidence that non-native earthworms occurred more frequently in areas with eastern redcedar-encroachment; instead, non-native earthworms were most likely to occur in tallgrass prairie. Our results suggest that, within prairies and woodlands of the Great Plains, native and non-native earthworms occur most frequently near roadways and in locations with moist soil. Because the few approaches for controlling invasive earthworms are only likely to be feasible on a small scale, findings from such local-scale studies are important for directing management to reduce earthworm impacts on biodiversity and ecosystem services.     

Pliocene Albizia (Fabaceae) from Jharkhand,eastern India: reappraisal of its biogeography during the Cenozoic in Southeast Asia

Albizia, a diverse tree genus, occupies monsoonal warm humid rain forests in tropical and subtropical regions. We recovered a well-preserved compound fossil leaf and two fossil fruits of Albizia (Fabaceae) from the latest Neogene (Rajdanda Formation: Pliocene) sediments of Jharkhand of the Chotanagpur Plateau, eastern India. On the basis of the architectural features of the fossil leaf, a new species is established as A. mahuadanrensis Hazra, Hazra and Khan, n. sp., characterised by a bipinnate, compound leaf having a rachis bearing opposite, asymmetrically ovate to sub-rhomboid leaflets, pulvinus on leaflet petiolule and brochiodromous secondary veins. Based on both morphological and anatomical characters of the fossil fruits, A. palaeoprocera Hazra, Hazra and Khan, n. sp. is erected, characterised by flattened to broadly linear shaped, wingless fruits; ovate-elliptic shaped seed chambers having ellipsoidal seeds in one series; irregularly polygonal to rectangular epidermal cells with oblique end walls and randomly oriented, scattered, paracytic stomata. Analysis of Albizia fossil occurrences indicates that the legume taxon was common in Neogene forests of India and elsewhere. The present-day distribution of the closely affiliated modern species of the fossil taxa indicates a warm and humid tropical environment during the time of deposition. We also review the biogeographic history of Albizia in India and other Asian countries.