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Hydrilla verticillata

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Lymnaea stagnalis

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Cornus amomum

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Misgurnus anguillicaudatus

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Pond Loaches live in rivers, ponds, streams, and other small bodies of water. Many live in low oxygen levels because fewer predators exist there.

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Eupatorium Perfoliatum; Asteraceae

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It thrives in average to wet soils with a consistent water source. It prefers full sun or part shade and tolerates both sandy and clay soil.

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Silky Dogwood prefers well-drained medium to acidic soils. They like to be grown in full sun, and partial shade and can tolerate full shade. More flowers will bloom with more sun.

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Agrimonia Parviflora

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Persicaria maculosa; Polygonaceae

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Grammotaulius nigropunctatus

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It prefers any fresh or brackish body of water. It is highly adaptable to a lot of growing conditions.

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The snail prefers stagnant to slow-moving waters with lush vegetation and likes nutrient-rich environments.

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Swamp Agrimony prefers full sun to light shade, moist conditions and can tolerate temporary flooding in the spring.

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The fly is found in temperate lakes, streams, and ponds. They are able to tolerate low oxygen concentrations and feed off of algae.

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Bombus

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Various bugs and bees visit the flowers on swamp agrimony. Birds often use the plant for their nests, as well.

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Silky Dogwood

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Common boneset

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Bumblebees tend to nest in relatively undisturbed areas such as shaded corners. They don't like prolonged exposure to the sun as this can heat up the nest too much.

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Redshank lives mostly on moist, rich soil. It likes cereals and other crops but isn't associated with one particular crop.

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Swamp Agrimony

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Redshank is rich in chlorophyll that helps plants get energy from the sun.

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Swamp Agrimony is native to Illinois and northeastern North America.

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Silky Dogwood is native to eastern and central parts of the United States as well as parts of Canada.

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It is native to the eastern USA and Canada and can be found in various kinds of wetlands.

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Genus Mentha

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Mint thrives in light soil with good drainage. They prefer a moist soil, rich with organic matter.

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Mint

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Bumblebee

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Mint is native to southern Europe and the Mediterranean region. It has been used by various peoples since at least 300 BC.

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Bumblebees are native to North America and they prefer higher altitudes or latitudes.

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Redshank

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https://mdc.mo.gov/discover-nature/field-guide/swamp-agrimony

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Description: This perennial wildflower is 2½-5' tall. The stout central stem is unbranched, terete, and light green, reddish green, or brownish green; it is covered with long hairs that are white or light brown. Along each stem, there are widely spreading alternate leaves. These leaves are odd-pinnate and up to 2' long and ½' across; each leaf has 9-17 primary leaflets and smaller secondary leaflets. The secondary leaflets are located between pairs of primary leaflets. Individual primary leaflets are 2-3" long and about one-third as much across; they are narrowly lanceolate, narrowly oblanceolate, or elliptic with wedge-shaped bottoms and acute tips. Leaflet margins are coarsely dentate. The upper surface of each leaflet is yellowish green and hairless, while the lower surface is short-pubescent. Secondary leaflets are similar to the primary leaflets, but they are much smaller in size (less than 1" long). Both the petiole and rachis of each compound leaf are pubescent; quite often, they have sparse long hairs. At the base of each leaf, there is a pair of large stipules that are fan-shaped and either coarsely dentate or cleft with pointed lobes.

The central stem terminates in a long spike-like raceme about ¾-2½' long. Robust plants also produce secondary racemes from the axils of the upper leaves that are shorter than the terminal raceme. These racemes are usually more or less erect, although longer racemes sometimes bend sideways to become nearly horizontal with the ground. The central stalk of the raceme is light green, terete, and short-pubescent. Numerous small flowers about ¼" across occur along the length of the raceme on short stalks about 1/8" long. Individual flowers consist of a tubular green calyx, 5 yellow petals, about 10 stamens, and a central pistil. The tubular calyx is turbinate in shape and 10-ribbed. The blooming period occurs from mid- to late summer and lasts about 1-2 months. Afterwards, the flowers are replaced by 1-2 seeded fruits about ¼" across. These small fruits have numerous hooked prickles along the upper rims of their persistent calyxes. Immature fruits are green, while mature fruits are brown. The root system is fibrous and rhizomatous. Clonal colonies of plants are often produced.

Cultivation: Swamp Agrimony prefers full sun to light shade, moist conditions, and loamy, silty, gravelly, or sandy soil. It tolerates temporary flooding during the spring.

Range & Habitat: Swamp Agrimony is occasional throughout Illinois (see Distribution Map), where it is native. Habitats include openings in floodplain woodlands, swamps, soggy thickets, gravelly seeps, riverbottom prairies and prairie swales, and roadside ditches. The preceding habitats can be either sandy or non-sandy.

Faunal Associations: The nectar and pollen of the flowers attract small bees and flower flies (Syrphidae). Larvae of the midge, Contarinia agrimoniae, feed on the flower buds, flowers, & developing seeds of Agrimonia spp. and other species in the Rose family, while an aphid, Macrosiphum agrimoniellum, sucks juices from the flowering stems. Insects that feed on the foliage of Agrimonia spp. include larvae of a sawfly (Fenella nigrita), larvae of a Gelechiid moth (Anacampsis agrimoniella), and larvae of a Tischeriid moth (Coptotriche agrimoniella). Because the foliage is bitter-tasting and high in tannins, it is usually avoided by mammalian herbivores. The bur-like fruits can cling to the fur of mammals, the feathers of birds, and the clothing of humans, which spreads the seeds to new locations.

Photographic Location: Near Cowle's Bog at the Indiana Dunes National Lakeshore in NW Indiana, where it was moist, partially shaded, and sandy.

Comments: Compared to other Agrimonia spp. in Illinois, Swamp Agrimony is easy to identify because its compound leaves have more primary leaflets (9-17) and its primary leaflets are more narrow in shape. To a greater extent than other species in this genus, Swamp Agrimony prefers habitats that are sunny and wet. Only a few flowers are in bloom at the same time; they are short-lived and not very showy because of their small size.

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Description: This perennial wildflower is 2½-5' tall. The stout central stem is unbranched, terete, and light green, reddish green, or brownish green; it is covered with long hairs that are white or light brown. Along each stem, there are widely spreading alternate leaves.

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Its blue berries are eaten by many songbirds and it is a host plant for the spring azure butterfly.

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Bees are extremely attracted to common bonesets because of their white flowers. It can also help humans as it reduces fever, can cause vomiting, and treat constipation.

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Caddis Fly

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Pond Leach

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Great Pond Snail

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Water Thyme

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Redshank likes the Great Lakes region and was introduced as an invasive species in North America. It likes to live on riverbanks and grows along the roadsides.

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Caddis flies are found worldwide but much prefer warmer conditions.

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Pond Loaches are native to eastern Asia and some parts of Africa, including Morocco and Ethiopia.

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It is native to parts of Europe, North America, and Asia (except its south region).

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It is native to the Indian Subcontinent, specifically Sri Lanka.

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Caddis flies are a food source for fish and they filter much of the algae and other organisms in the water.

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Pond Loaches are bottom-dwelling scavengers that feed off of organic matter like algae. Some people in eastern Asia have eaten these fish.

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The snails are great scavengers and will eat any leftover or dead carcass.

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Water Thyme is technically a producer in the food web. It also provides cover for predators and prey.

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Mint provides food for various insects and is an important food source for bees and other pollinators.

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Bees are one of the most important pollinators in our food chain. They visit flowers, carrying the pollen from the male part of flowers to the female part, allowing fertilization.

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https://storymaps.arcgis.com/stories/466f017e72a2465a8f0867bacbb402d6#:~:text=Caddisflies%20are%20extremely%20important%20to,cause%20taste%20and%20odor%20problems.

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Trichoptera (caddisflies)

A caddisfly assemblage can provide information on the local aquatic habitats, based on their biological and distributional information. As such, they are of use in both Quaternary and forensic science. Caddisflies have been found on submerged bodies ( Wallace et al., 2008) and used to determine the duration of time since submergence. The value of the particular families of caddisflies varies with the circumstance ( Figure 3).

Net-spinning caddisflies (Hydropsychidae), although of value in Quaternary studies, do not necessarily provide a specific means of determining the postmortem submerged interval. Those species of caddisfly that make their cases from vegetable matter and sand and other fragments can often be more useful. For example, Wallace et al. (2008) used the seasonality and the duration and timing of life stages, including the onset of dormancy, of two species of Limnephilid caddisflies to estimate the time of submergence of a body at a crime scene. These forensic entomologists were able to infer that the body was submerged between late April and late May.

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l. Phryganeidae (10 Genera, 28 Species)

These large caddisflies have distinctive cases made mostly of pieces of vegetation that are spirally wound or in concentric rings. When disturbed, they readily abandon their cases, but may re-enter them. Larvae are distinctive, generally having a boldly striped head that is more prognathous than other Integripalpia (Fig. 65). They also have a prominent prosternal horn and lack significant sclerotization of the mesonotum. Identification of larvae to species is not possible in some genera. Larvae may be found among vegetation and detritus along streams of all sizes, in marshes, in temporary and permanent ponds, and even in lakes where they may occur far from shore. While many are mostly predators, vegetation is also consumed, especially by early instars. Life cycles are probably 1 year.

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# data[].attributes.attachment_link.description
Read full chapter URL: https://www.sciencedirect.com/science/article/pii/B9780123814265000259 A caddisfly assemblage can provide information on the local aquatic habitats, based on their biological and distributional information. As such, they are of use in both Quaternary and forensic science. Caddisflies have been found on submerged bodies ( Wallace et al., 2008) and used to determine the duration of time since submergence.

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Loach - Description, Habitat, Image, Diet, and Interesting Facts

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Everything you should know about the Loach. The Loach is an elongated eel-shaped fish, with fleshy, whisker-like appendages near it's mouth.

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https://evodevojournal.biomedcentral.com/articles/10.1186/s13227-020-00169-4#:~:text=Lymnaea%20stagnalis%20is%20a%20freshwater,most%20southern%20region%20%5B4%5D.

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Stuart RF King, Reviewing Editor and Peter Rodgers, Senior Editor

Stuart RF King, eLife, United Kingdom;

Abstract

Only a limited number of animal species lend themselves to becoming model organisms in multiple biological disciplines: one of these is the great pond snail, Lymnaea stagnalis. Extensively used since the 1970s to study fundamental mechanisms in neurobiology, the value of this freshwater snail has been also recognised in fields as diverse as host-parasite interactions, ecotoxicology, evolution, genome editing and 'omics', and human disease modelling. While there is knowledge about the natural history of this species, what is currently lacking is an integration of findings from the laboratory and the field. With this in mind, this article aims to summarise the applicability of L. stagnalis and points out that this multipurpose model organism is an excellent, contemporary choice for addressing a large range of different biological questions, problems and phenomena.

Introduction

In ancient Greece, over 2,400 years ago, it was already recognised that by studying animals we could learn much about ourselves. Over the centuries since then, it has become clearer that some species are highly suitable in the fields of medical, basic and applied biological research ( Ericsson et al., 2013). However, when considered carefully, there is perhaps only a limited set of animal species that are versatile enough to lend themselves to become model organisms in multiple biological disciplines ( Frézal and Félix, 2015; Hilgers and Schwarzer, 2019; Markow, 2015; Phifer-Rixey and Nachman, 2015).

In the second half of the 20 th century, one booming line of research has focused on molluscs. Neuroscientists such as the Nobel Prize winners Alan Hodgkin, Andrew Huxley and Eric Kandel recognised these animals' potential as models for understanding basic neurobiological processes ( Hodgkin and Huxley, 1952; Kupfermann and Kandel, 1969; Wachtel and Kandel, 1967). One particularly well-suited mollusc for this type of research is the freshwater pond snail, Lymnaea stagnalis, which has been used extensively since the 1970s to study the functioning of the nervous system from molecular signalling to behaviour.

The value of L. stagnalis also has been recognised in a wide range of applied biological fields. These include the study of host-parasite interactions, ecotoxicology, evolution, developmental biology, genome editing, 'omics' and human disease modelling. This extensive suitability stems from the most obvious advantages of L. stagnalis: its well-known anatomy, development (both of the embryonic and post-embryonic processes), and reproduction biology; its well-characterised central and peripheral nervous and neuroendocrine systems from key molecules to behavioural processes; and its readily accessible and mostly large neurons. There is also a growing body of available sequence data with an impending annotated genome and the option to use new technical approaches such as genome editing. Taking all of the above into consideration, these advantages simplify the study of different scientific topics integrated from the molecular to the population level.

This article is a tribute to over 50 years of research with L. stagnalis that has resulted in a considerable contribution to the understanding of general biological processes. Here, we present the essential background information on the natural history of this freshwater snail. We also provide an overview of the ground-breaking and recent information on different research fields using L. stagnalis (and snails in general). Our aim is to showcase L. stagnalis as a contemporary choice for addressing a wide range of biological questions, problems and phenomena, to inspire more researchers to use this invertebrate as a model organism, and to highlight how findings from the laboratory and the field could be better integrated.

Natural history of L. stagnalis

Initially described by Linnaeus in 1758 as Helix stagnalis, the species now known as L. stagnalis is generally referred to as the great pond snail (Panpulmonata; Hygrophila; Lymnaeidae). It is found throughout Northern America, Europe, and parts of Asia and Australia ( Atli and Grosell, 2016; Zhang et al., 2018a; Figure 1 ). The snails inhabit stagnant and slowly running shallow waters rich in vegetation and are mainly herbivores, preferring algae, water plants and detritus ( Lance et al., 2006). They are active all year round (even when there is a layer of ice on the water) but typically reproduce from spring to late autumn ( Nakadera et al., 2015). They do not have a clear day-night rhythm, but display sleep-like behaviour ( Stephenson and Lewis, 2011) and are more likely to lay eggs during daytime ( Ter Maat et al., 2012). They are light to dark brown in colour and relatively large for pond snail species, with their spiral shells reaching lengths of up to 55 mm ( Benjamin, 2008). In highly oxygenated water, they absorb oxygen directly across their body wall; but when dissolved oxygen levels drop, they switch to breathing via a lung accessed by a respiratory orifice called the pneumostome ( Lukowiak et al., 1996).

L. stagnalis serves as the intermediate host for parasites including flatworms responsible for diseases such as fascioliasis (liver fluke and river rot) and cercarial dermatitis (swimmer's itch) in humans ( Adema et al., 1994; Davison and Blaxter, 2005; Ferté et al., 2005; Núñez et al., 1994; Skála et al., 2020). Laboratory and field studies showed that penetration of a parasite into a snail will initiate a chronic infection in which the parasite alters snail neurophysiology, metabolism, immunity, growth and reproduction ( Kryukova et al., 2014; Langeloh and Seppälä, 2018; Vorontsova et al., 2019). These studies have also investigated how selection acts on immune defence traits ( Langeloh et al., 2017). Investigations of the natural history of L. stagnalis, which focus on host-parasite associations, aid the development of novel control measures that reduce snail-mediated parasitic transmissions. Primary predators of juveniles and adults include leeches, crayfish and fish, some of which snails can detect via chemicals that the predators emit ( Dalesman and Lukowiak, 2012).

The life cycle and reproductive biology of the species are well-characterised ( Ivashkin et al., 2015; Koene, 2010; Mescheryakov, 1990; Morrill, 1982; Figure 2A ). Embryos develop inside transparent eggs packaged in a translucent gelatinous mass allowing observers to follow their developmental stages in detail over the 11-12 days to hatching. The time from laid eggs to mature reproductive adults can be as short as two months depending on the temperature, photoperiod, feeding regime and mate availability at the location where they are being raised. In their natural habitat, they have been found to reach an age in excess of one year, but in the laboratory they live longer, up to two years ( Janse et al., 1988; Nakadera et al., 2015). For laboratory breeding, a large and genetically diverse breeding stock is recommended as this will facilitate a well-standardised stock population without too much inbreeding. The largest and longest-maintained breeding facility is found at the Vrije Universiteit in Amsterdam, where L. stagnalis has been bred continuously for over 50 years ( Nakadera et al., 2014).

Its well-characterised embryonic and post-embryonic processes have promoted extensive use of L. stagnalis in the field of developmental biology. This snail has helped us to understand the mechanisms underlying shell formation ( Hohagen and Jackson, 2013), the transfer of non-genetic information to the developing embryos ( Ivashkin et al., 2015), and resource allocation during development ( Koene and Ter Maat, 2004). Moreover, studies with L. stagnalis has also helped develop and evaluate models in physiology, such as the "dynamic energy budget" model ( Zonneveld and Kooijman, 1989; Zimmer et al., 2014).

This snail is a simultaneous hermaphrodite, meaning that mature individuals express a functional male and female reproductive system at the same time within one body. Despite having two functional sexes, specimens copulate unilaterally; one individual plays the male role and the other the female role within one mating interaction ( Hoffer et al., 2017). There is no obligate alternation of sexual roles, but when both individuals of a mating pair are motivated to mate in the male role they can perform a second copulation with the same partner in the opposite sexual role ( Koene, 2010; Ter Maat et al., 2012; Van Duivenboden and Maat, 1985). An integrated laboratory and field study showed that, in the wild, most individuals are born during the spring and summer seasons and generations partly overlap because the latter cohorts overwinter and overlap with mature individuals of the new spring cohort ( Figure 2B ; Nakadera et al., 2015). The same study showed that both age and size significantly affected the sex role decision under laboratory conditions. This species is quite fecund in the laboratory: snails from the mass culture in Amsterdam produce a large number of offspring all year round ( Nakadera et al., 2014); however, an initial field study found a more moderate fecundity rate in natural populations ( Nakadera et al., 2017). In the laboratory, specimens produce on average 2-3 egg masses per week each containing 100-150 eggs, depending on the body size of the individual ( Nakadera et al., 2014). The hatching rate under laboratory conditions is generally above 90% ( Hoffer et al., 2017). Based on laboratory, semi-field and field studies, explicit inbreeding or self-fertilisation depression for this species have been found to be absent ( Coutellec and Lagadic, 2006; Escobar et al., 2011; Koene et al., 2008; Puurtinen et al., 2007) or very unlikely ( Coutellec and Caquet, 2011), however the reasons for this remain unclear ( Box 1). Nevertheless, eggs are preferentially outcrossed with sperm from mating partners, which can be stored for two months, and individuals only use their own 'autosperm' when this 'allosperm' is not available ( Nakadera et al., 2014).

Box 1.

Why is inbreeding depression less strong in L. stagnalis than in related freshwater snail species?

How different are long-term laboratory-bred strains from natural populations as a result of different selection pressures influencing development, mating propensity, self-fertilisation, learning and/or changes in sensitivity due to changing biotic and abiotic factors?

How can the knowledge about host-parasite interaction be applied to control the spread of parasites in the natural habitat?

How phenotypically plastic or evolutionarily adaptable is this species to changes in biotic and abiotic conditions in its habitat (e.g., temperature, light and/or chemical pollution, and resulting changes in ecosystem composition)?

Why are sinistral individuals not found more often in natural populations and what does that mean for the natural selection pressures on this chiral morph?

Are the detection and avoidance of positive and negative stimuli only present in the laboratory or is this learned behaviour also exhibited under field conditions (e.g., predicting presence of food, mating partners and/or predators)?

How can the knowledge about the regulatory mechanisms underlying reproduction be better used to understand the evolution and flexibility of the hermaphroditic lifestyle?

A gold standard model organism for neuroscience

The squid Loligo forbesii and sea hare Aplysia californica were the first molluscan models for examining neuronal processes. L. stagnalis emerged shortly afterwards, and was described as "a reductionistic, yet sophisticated model to address fundamental questions in learning and memory" ( Rivi et al., 2020). Molluscs were used extensively in the field of neurobiology in the 20th century, typically because their central nervous systems were, in most cases, more accessible than those of vertebrate animals. Technical developments since then mean many such experiments can now be performed on vertebrates as well, yet we would argue that invertebrates still have substantial advantages for our understanding of the central nervous system.

The relatively simple central nervous system of L. stagnalis is organised in a ring of 11 interconnected ganglia ( Figure 3A,B ) with ~25,000 neurons. The neurons are mostly large (30-150 µm in diameter) and their bright, orange-coloured cell bodies are located on the surface of the ganglia ( Figure 3B ; Kemenes and Benjamin, 2009). Thus they are readily accessible for experimental purposes, simplifying investigations of neural clusters, circuits and even single neurons, which can be reliably identified for functional examination using a variety of approaches such as electrophysiological, molecular and analytical techniques ( Crossley et al., 2018; de Hoog et al., 2019; El Filali et al., 2015; Harris et al., 2012; Kemenes et al., 2011; Lu et al., 2016; Samu et al., 2012; Wagatsuma et al., 2005; Zhang et al., 2018b).

Individual neurons ( Figure 3C ; Benjamin and Crossley, 2020) and their synaptic connectivity were identified as parts of circuits controlling behaviours ( Audesirk et al., 1985; Benjamin, 2012; McCrohan and Benjamin, 1980a; McCrohan and Benjamin, 1980b; Syed and Winlow, 1991; Syed et al., 1990). Combining this knowledge with an understanding of the molecular mechanisms, often from laboratory studies, has helped produce an integrated picture of the processes underlying learning and memory, such as consolidation, reconsolidation, extinction and forgetting. The molecular pathways involved in memory formation in L. stagnalis were recently identified, providing further evidence the mechanisms of learning and memory consolidation are conserved across phylogenetic groups in a variety of learning paradigms, including non-associative or associative learning, and operant or classical conditioning ( Benjamin and Kemenes, 2013; Fulton et al., 2005; Josselyn and Nguyen, 2005; Kemenes and Benjamin, 2009; Kemenes et al., 2002; Marra et al., 2013; Michel et al., 2008; Nikitin et al., 2008; Park et al., 1998; Pirger et al., 2010; Pirger et al., 2014a; Pirger et al., 2014b; Ribeiro et al., 2003; Rivi et al., 2020; Sadamoto et al., 1998; Sadamoto et al., 2010; Schacher et al., 1988; Vigil and Giese, 2018; Wan et al., 2010). Recently studies have also revealed differences in learning ability at the behavioural level between situations in the laboratory and the field (e.g., Dalesman and Lukowiak, 2012; Dalesman et al., 2015; Dalesman, 2018).

The well-characterised proximate processes at the molecular, cellular and circuit levels mean studying this simple nervous system has the potential to provide insights into how snails can respond appropriately to environmental challenges (e.g., climatic change or pharmacologically active compounds). Also, since their behaviours are generated by reflexive and central pattern generator networks similar to those of vertebrates ( Katz and Hooper, 2007), results from snails offer insights into the fundamental processes important for these animals too.

Finally, recent developments have enabled this species to be used as a model for understanding the basis of neurodegenerative diseases. Comparative analyses have yielded several homologs to human genes linked to ageing and neurodegenerative diseases in A. californica and this species has proved well suited for studying these processes ( Moroz et al., 2006; Moroz and Kohn, 2010). Similar molecular sequences have been identified in L. stagnalis ( Fodor et al., 2020b). With the appropriate genetic background, its accessible central nervous system and relatively long and well-characterised life span mean L. stagnalis is highly suitable for studying the biological mechanisms of ageing, age-related memory loss and neurodegenerative diseases, such as Parkinson's and Alzheimer's diseases ( Arundell et al., 2006; de Weerd et al., 2017; Ford et al., 2017; Hermann et al., 2007; Hermann et al., 2020; Maasz et al., 2017; Patel et al., 2006; Pirger et al., 2014b; Scutt et al., 2015; Vehovszky et al., 2007; Yeoman and Faragher, 2001; Yeoman et al., 2008).

Ecotoxicology and risk assessment in a changing global environment

It has become clear that in the globalised world, climate change, light pollution, micro- and nanoplastics, and pharmacologically active compounds all pose a challenge to animal life. These challenges affect the availability of suitable habitats and reduce the quality of the land, lakes and rivers. They also change the environmental composition of pathogens, parasites, competitors and invaders. Understanding how global ecosystems are adapting to pollution is a complex problem; it requires researchers to monitor natural populations and conduct laboratory studies to discover the bases of adaptations or the lack thereof ( Markow, 2015).

Molluscs the second most diverse animal group and considered to be excellent indicators of ecosystem health. For example, L. stagnalis is a sensitive and reliable species for such studies ( Amorim et al., 2019), in a large part because of its well-characterised developmental and reproductive biology as described above. The major targets in this field of study have been metal-risk assessment ( Crémazy et al., 2018; Pyatt et al., 1997; Vlaeminck et al., 2019), the effects of pesticides ( Coutellec et al., 2008; Lance et al., 2016; Tufi et al., 2015; Vehovszky et al., 2015), nanotoxicology ( Hudson et al., 2019; Stoiber et al., 2015), the development of toxicokinetic models ( Baudrot et al., 2018), immunocompetence analyses ( Boisseaux et al., 2018; Gust et al., 2013b), and global warming risk assessment ( Leicht et al., 2017; Leicht and Seppälä, 2019; Teskey et al., 2012). Studies on L. stagnalis have measured toxicological values such as mortality concentrations (e.g., LC50) and impairment of reproduction (e.g., EC50) but also sub-lethal and more sensitive endpoints such as reproductive success, growth, cellular and molecular biomarkers that may be coupled with behavioural responses ( Amorim et al., 2019).

L. stagnalis has also been recognised as a useful organism to examine the effects of pharmacologically active compounds and micro- and nanoplastics on aquatic organisms ( Amorim et al., 2019; Charles et al., 2016; Ducrot et al., 2014; Gust et al., 2013a; Horton et al., 2020; Pirger et al., 2018; Zrinyi et al., 2017). However, it must be pointed out that researchers need to be critical of the generalisability of results while performing such experiments since there are differences between the endocrine system of molluscs and vertebrates; molluscs, for example, do not have functional oestrogen receptors ( Eick and Thornton, 2011; Lagadic et al., 2007; Scott, 2012). It is also important to recognise that molluscs are not suitable for some types of ecotoxicological studies and they cannot always substitute for fish.

Notably, L. stagnalis is the first aquatic non-arthropod invertebrate model organism to be recognised in environmental risk assessments. The developed standard reproduction test was officially approved by the national coordinators of the Organisation for Economic Cooperation and Development ( OECD, 2016) thus paving the way for investigating ecotoxicological effects in more detail. Such information will contribute to a more complete picture of the mode of action of potentially toxic substances and other environmental factors and provide assessments of risk for individual species of different types and wider ecosystems.

Combining evolution and natural history

Within the field of evolutionary biology, L. stagnalis has helped us to understand the evolution of several phenomena. Left-right asymmetry is a general evolutionary phenomenon seen across a variety of species, including humans where the congenital condition situs inversus results in the mirrored position and shape of the heart and liver ( Blum et al., 2014; Oliverio et al., 2010; Palmer, 2009; Palmer, 2016). The coil or chirality of snail shells is one of the more spectacular outward manifestations of this asymmetry. Snails found in nature can have shells that coil either to the right or left, with most species being right-coiling. Specimens of L. stagnalis that coil in the opposite left-winding, or sinistral, direction are rare and often categorised as 'unlucky' because their different chirality makes it difficult for them to mate the more usual, right-winding individuals ( Davison et al., 2009). Left-winding snails are also less able to learn in a mate-choice context ( Koene and Cosijn, 2012). The existence of the two different morphs has made this species ideal for studying chiromorphogenesis, i.e. the first step of left-right symmetry breaking. Genes and signalling pathways that are responsible for snail coiling have been identified ( Abe et al., 2014; Davison et al., 2016; Kuroda, 2014; Kuroda, 2015; Kuroda et al., 2016), and similar signalling pathways are required for vertebrate chiromorphogenesis as well ( Kuroda, 2015). Studies on L. stagnalis can give important insights into the evolution of body plans in other phyla, and may have wider medical implications, including an understanding of situs inversus.

L. stagnalis has also played a crucial role in studies into the evolution of hermaphroditism and its consequences for sexual selection. This area of research relies heavily on a solid understanding of the natural history of this species. Selection of sexual traits that affect mating success was previously considered not to act in simultaneous hermaphrodites ( Charnov, 1979; Darwin, 1871; Greeff and Michiels, 1999; Morgan, 1994). However, recent research, including work with L. stagnalis, has contradicted this earlier conclusion ( Anthes et al., 2010; Baur, 1998; Chase, 2007; Janicke et al., 2016; Michiels, 1998; Nakadera and Koene, 2013). Unilateral mating of L. stagnalis offered a unique opportunity to test whether sexual selection acts independently on the two sexual roles of a simultaneous hermaphrodite ( Anthes et al., 2010; Arnold, 1994; Hoffer et al., 2017). Recent experiments have also revealed that male and female reproductive strategies can be optimised independently in this species. This was done by measuring sexual selection gradients (also called Bateman gradients), which reveal the relationship between the number of matings and the reproductive success of the sexual functions ( Anthes et al., 2010). Experiments with L. stagnalis showed that this mating system seems largely male-driven, and that the sexual selection gradients are consistently positive for the male function but change over time to benefit the opposite sex ( Anthes et al., 2010; Hoffer et al., 2017; Janicke et al., 2016; Pélissié et al., 2012). These pioneering works, which measured and quantified the processes of sexual selection and their underlying mechanisms, thus incorporated this hermaphrodite into the general Darwin-Bateman paradigm that had so far mainly been tested on separate-sexed species. They also described both the evolutionary potential and limitations of hermaphrodite animals and revealed important practical applications for the conservation of wildlife.

New opportunities from a growing multi-omics coverage

From about 1980, continued attention was given to the physiological characterisation of L. stagnalis, but more recent research has focussed on an 'omics' approach to better understand the underlying molecular processes ( Santama et al., 1993; Santama et al., 1995a; Santama et al., 1995b; Santama and Benjamin, 2000). Due to its pre-eminence as a model system in neuroscience, early molecular studies tended to focus on the central nervous system ( Feng et al., 2009; Johnson and Davison, 2019). The favourable anatomical features enabled the accumulation of peptidomic data from the mass spectrometry of single neurons ( Perry et al., 1999; Worster et al., 1998), making the neuropeptidergic system the most intensely studied part of the central nervous system ( Buckett et al., 1990; Perry et al., 1998). Taking advantage of a variety of platforms available for nucleotide sequencing: Sanger ( Davison and Blaxter, 2005; Sadamoto et al., 2004; Swart et al., 2019), Illumina ( Korneev et al., 2018; Sadamoto et al., 2012; Stewart et al., 2016), BGISEQ ( Jehn et al., 2018) and Oxford Nanopore ( Fodor et al., 2020a), many sequencing methodologies have been successfully applied to this species.

Extensive genomic, transcriptomic and peptidomic data for L. stagnalis are available in the NCBI database. Four major transcriptome datasets were established by sequencing mRNA from the central nervous system ( Bouétard et al., 2012; Davison and Blaxter, 2005; Feng et al., 2009; Sadamoto et al., 2012), and then used to identify genes and proteins, thus providing a solid genetic background for L. stagnalis. Furthermore, an unannotated draft genome is already available and a collaborative effort is underway to produce an annotated genome ( Johnson and Davison, 2019) which would largely solve the problem of the lack of molecular information that has so far inhibited research in the L. stagnalis model system ( Rivi et al., 2020). Approximately 100 (neuro)peptides have been identified so far ( Benjamin and Kemenes, 2020), encoded by genes involved in various regulatory processes ( Table 1 ). These findings contributed to a global understanding of the natural history of L. stagnalis by characterising the molecular and cellular processes underlying chirality, reproduction, immune processes, host-parasite interaction, and acute and chronic adaptive responses to toxic substances in the environment.

Table 1.

Molecule

Abbreviation

Function

Accession number

Reference

actin-related diaphanous genes (1, 2)

dia 1, dia 2

development, chirality

KX387869, KX387870
KX387871, KX387872
Kuroda et al., 2016

insulin-related peptides
(I, II, III, V, VII)

MIPs

development

CAA41989; P25289; AAB28954; AAA09966; AAB46831

Smit et al., 1991; Smit et al., 1992; Smit et al., 1993b; Smit et al., 1996; Smit et al., 1998

pituitary adenylate cyclase-activating polypeptide-like molecule

PACAP-like

learning and memory

-
Pirger et al., 2010

cAMP response element-binding proteins (1, 2)

CREB 1
CREB 2

learning and memory

AB041522; AB083656
Sadamoto et al., 2004

glutathione reductase and peroxidase

Gred
Gpx

metabolic detoxification

FJ418794,
FJ418796
Bouétard et al., 2014

Furthermore, the CRISPR/Cas9 genome editing method has recently been applied to molluscs ( Henry and Lyons, 2016; Perry and Henry, 2015). In L. stagnalis, it was used to knock out the gene responsible for coiling direction during development, leading to a better understanding of chirality in the life of the two morphs ( Abe and Kuroda, 2019). The establishment of genome editing in L. stagnalis opens up significant opportunities for functional genomics to investigate the role of specific genes, for example, in snail developmental, toxicology and immunobiological studies.

Conclusion

Research on model organisms has been essential to developing the current understanding of how life works. The unique features of L. stagnalis make it an excellent experimental system to complement the classic invertebrate ( C. elegans, D. melanogaster) and vertebrate ( D. rerio, M. musculus) models. Research utilising this species is expected to lead to future breakthroughs in a number of scientific fields, especially in neuroscience and evolutionary biology. For example, as a simultaneously hermaphroditic outcrossing species, it presents the opportunity to test the generality of hypotheses that are mainly based on non-hermaphroditic or self-fertilising models. There is considerable information about the natural history of L. stagnalis compared to some other model species, but we feel some areas of research using L. stagnalis - in particular neurobiology and ecotoxicology - would benefit by extending more of their studies out of the laboratory and into the field. We believe that a deeper integration of information from field studies with input from laboratory findings - such as applying experimental designs and approaches developed in the laboratory to populations in the wild - will provide future opportunities for further innovation ( Box 2). Such efforts could address the unanswered questions regarding this model organism (see Box 1). Significantly, emerging recent technical approaches such as pocket-sized sequencing devices, especially with their impending breakthrough also in protein sequencing, start allowing researchers to perform more experiments in the field such as following molecular mechanisms of learning.

Box 2.

It needs to be established at what level L. stagnalis can function as a model for medical research such as neurodegenerative disease and be a substitute for standard vertebrate models. This requires a better understanding of how such functions affect this species in its natural habitat.

The new molecular techniques and available 'omics' data provide an incentive for research that aims to understand the mechanisms underlying natural history processes such as sex allocation, simultaneous hermaphroditism, reproductive success, chirality and learning.

The knowledge about learning and decision-making in the laboratory needs to be extended to field populations to promote future developments in, for example, neural network-inspired robotics.

Biographies

István Fodor is in the NAP Adaptive Neuroethology, Department of Experimental Zoology at the Balaton Limnological Institute, Centre for Ecological Research, Tihany, Hungary

Ahmed AA Hussein is in the Department of Ecological Sciences, Faculty of Sciences at Vrije Universiteit, Amsterdam, the Netherlands

Paul R Benjamin is at Sussex Neuroscience, School of Life Sciences, University of Sussex, Brighton, United Kingdom

Joris M Koene is in the Department of Ecological Sciences, Faculty of Sciences at Vrije Universiteit, Amsterdam, the Netherlands

Zsolt Pirger is in the NAP Adaptive Neuroethology, Department of Experimental Zoology at the Balaton Limnological Institute, Centre for Ecological Research, Tihany, Hungary

Funding Statement

The funders had no role in study design, data collection and interpretation, or the decision to submit the work for publication.

Contributor Information

Stuart RF King, eLife, United Kingdom.

Peter Rodgers, eLife, United Kingdom.

Funding Information

This paper was supported by the following grant:

National Brain Research Project 2017-1.2.1-NKP-2017-00002 to Zsolt Pirger.

Additional information

Competing interests

No competing interests declared.

Author contributions

Conceptualization, Writing - original draft, Writing - review and editing.

Conceptualization, Writing - original draft.

Conceptualization, Writing - original draft.

Conceptualization, Visualization, Writing - original draft.

Conceptualization, Supervision, Funding acquisition, Visualization, Writing - original draft, Writing - review and editing.

Data availability

All data generated during the preparation of this review are included in the manuscript.

References

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The unlimited potential of the great pond snail, Lymnaea stagnalis

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Only a limited number of animal species lend themselves to becoming model organisms in multiple biological disciplines: one of these is the great pond snail, Lymnaea stagnalis. Extensively used since the 1970s to study fundamental mechanisms in neurobiology, ...

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https://nas.er.usgs.gov/queries/FactSheet.aspx?SpeciesID=6#:~:text=Size%3A%20Stems%20grow%20up%20to,Sri%20Lanka%20(Schmitz%20et%20al.

http://erie.cce.cornell.edu/invasive-species/ecological-impacts-of-hydrilla#:~:text=Although%20hydrilla%20can%20provide%20habitat,additional%20cover%20provided%20by%20hydrilla.

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Species Overview

Native to: India ( Hydrilla verticillata's dioecious type originates from southern India. Hydrilla's monoecious type is probably from Korea)

Hydrilla was introduced into Florida water bodies in 1950-1951.It was thought to have been introduced to the Tampa and Miami areas as an aquarium plant. By the 1970s, it was established throughout Florida waters and in most drainage basins. Hydrilla can grow to the surface of waters as deep as 25ft and form dense mats and can still be found in all types of water bodies.

Species Characteristics

Family: Hydrocharitaceae

Habit: submersed aquatic, profusely branched, herbaceous perennial with stems up to 20 ft long; obligate wetland plant

Leaves: smallstrap-like, pointed; whorled, saw-toothed

Flowers: (Florida biotype) female flowers only; solitary, tiny, white, float on the surface; threadlike stalks attached at leaf axils near the stem tips

Reproductive parts: turions ("buds" in some of the leaf axils), dark green, cylindrical, to 1/4 in. round. Subterranean turions ("tubers"); yellowish, potato-like, attached to the root tips in the hydrosoil, to 1/2 in. long, 1/2 inch broad; readily reproduces by fragments; single tuber can grow to produce more than 6,000 new tubers per square m

Seeds: Florida biotype does not produce

Distribution in Florida: statewide

Impacts

Hydrilla has widescale impacts in Florida waters and is highly adaptable to a variety of growing conditions. It can grow in almost any freshwater system including springs, lakes, marshes, ditches, rivers and tidal zones. Hydrilla can grow in water as shallow as a few inches and up to 20 feet deep. It can grow in as little as 1% of full sunlight.

Hydrilla continues to be sold through aquarium supply dealers and over the internet, despite being a Federal Noxious Weed and a Florida Prohibited Aquatic Plant. Each stem on a Hydrilla plant can grow 1-4 inches per day. Therefore, when hydrilla invades water bodies, ecologically-important native submersed plants such as pondweeds ( Potamogeton spp.), tapegrass ( Vallisneria americana) and coontail ( Ceratophyllum demersum) are shaded out by hydrilla's thick mats, or are simply outcompeted and eliminated.

Each year in Florida, millions of dollars are spent on herbicides and mechanical harvesters in an effort to place hydrilla under "maintenance control." Without management, hydrilla slows water flow and clogs irrigation and flood-control canals and interferes with boating (both recreational and commercial) and prevents swimming and fishing. Dense infestations can alter water chemistry and dissolved oxygen levels.

Hydrilla is a prohibited plant and therefore, not recommended by UF/IFAS. Hydrilla is a prohibited plant according to the USDA Noxious Weed List and the Florida Prohibited Plant List. The UF/IFAS Assessment lists Hydrilla as prohibited. It is listed by FLEPPC as a Category l invasive species due to its ability to invade and displace native plant communities.
Preventive Measures
Avoid introducing hydrilla into water bodies. Use best practices to prevent introduction by cleaning boat trailers, propellors, diver gear and live bait wells. Transporting plant fragments on boats, trailers, and in livewells is the main source of introduction into new lakes and rivers. Do not use hydrilla in aquariums or ornamental ponds. Opt for native submersed aquatic plants such as, sago Pondweed ( Potamogeton pectinatus), bladderwort ( Utricularia floridana), coontail ( Ceratophyllum demersum), fanwort ( Cabomba caroliniana) or southern naiad ( Najas guadalupensis).
Cultural/Physical
Small infestations of Hydrilla may be removed either manually or using hand tools, such as a rake. In some cases, lake drawdowns may help manage hydrilla by letting the exposed plants die and decompose.
Mechanical
Mechanical harvestors can be used toremove hydrilla from the water and transport it to shore for disposal. One drawback in the use of mechanical harvesters is that cuttings of hydrilla, which are not removed from the water, help to spread this weed.
Biological
Currently, four insects and one fish have been released to control hydrilla, but only two of these insects are established, and only one is commonly associated with hydrilla in the southeastern U.S. Click here to learn more.

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| Center for Aquatic and Invasive Plants | University of Florida, IFAS

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Native to: India ( Hydrilla verticillata's dioecious type originates from southern India. Hydrilla's monoecious type is probably from Korea) Hydrilla was introduced into Florida water bodies in 1950-1951.It was thought to have been introduced to the Tampa and Miami areas as an aquarium plant.

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Plant DetailShow Menu

Cornus amomum is often confused with:

Plants that fill a similar niche:

Phonetic Spelling

KOR-nus ah-MOH-mum

Description

Silky dogwood is a native large to a medium-sized deciduous shrub in the Cornaceae (dogwood) family. It is native to central and the eastern part of the United States and parts of Canada and is typically found in swamp borders, wetlands, and near streams and ponds. This shrub is multi-stemmed, open, and rounded and may grow 6 to 12 feet tall.

Clusters of creamy white flowers bloom in the spring followed by blue drupes that ripen late summer. The leaves and twigs have silky hairs. A distinctive feature of this shrub is the dark brown pith of 1 to 2-year-old stems and dark-reddish twigs.

The genus name, Cornus, is Latin from the word, cornu, which means "horn." This refers to the hardness of the wood. This shrub has several common names including swamp dogwood. This references its usual habitat.

The silky dogwood prefers well-drained medium to wet acidic soils. It may be grown in full sun, partial shade, and even tolerates close to full shade. The roots need to be kept cool and moist during the summer. It is recommended that 2 to 4 inches of mulch be added to protect the plant's roots. If the shrub branches touch the ground they may form roots at the nodes. Thickets may result if the growth of the shrub is not controlled. The shrub also may be propagated by seeds, cuttings, or layering.

The creamy white clusters of blooms appear at the tips of the branches from May to June. The leaves are dark green, alternate, simple, pubescent, and acutely veined with a smooth margin. The berry-like drupes appear in drooping clusters from August to September and attract many birds. The twigs are usually dark reddish-purple with fine hairs. The older bark is smooth and gray.

This is a good shrub selection for moist to wet areas of the landscape. It is not overly ornamental and is somewhat wild and unkempt for placement in prominent areas. Consider the silky dogwood for moist woodlands, naturalized areas, or sites near streams/ponds. It is also effective for erosion control.
NC Native Shrub Seasons of interest:
Bloom: Spring Fruit: Summer Twigs: Winter
Quick ID Hints:

open, rounded, medium-sized shrub

silky hairs on the leaves and twigs

creamy white clusters of flowers in the spring

drupes ripen to dark blue in late summer

dark-reddish purple twigs

smooth gray bark

dark brown pith in 1 to 2-year-old stems

Insects, Diseases, and Other Plant Problems: The silky dogwood has no serious insects or disease problems. It may be susceptible to scale. Borers and leaf miners are potential pests. Leaf spot, crown canker, blights, root rot, and powdery mildew may occur. Thickets may form if the shrub is not maintained.

See this plant in the following landscape:

Cultivars / Varieties:

'Indigo'

Tags:

#showy flowers#deciduous#rain garden#deciduous shrub#nectar plant#native shrub#riparian#wetlands#erosion control#showy fruits#stream banks#pond margins#specialized bees#cpp#fire medium flammability#NC native#cream flowers#pond garden#native garden#acidic soils tolerant#pollinator plant#naturalized area#nesting sites#larval host plant#food source summer#food source fall#food source herbage#food source nectar#food source pollen#Coastal FACW#Piedmont Mountains FACW#wet soils tolerant#bird friendly#food source soft mast fruit#butterfly friendly#bee friendly#black walnut toxicity tolerant#Audubon#woodland garden#hedge#stormwater garden#wildlife friendly

Cultivars / Varieties:

'Indigo'

Tags:

#showy flowers#deciduous#rain garden#deciduous shrub#nectar plant#native shrub#riparian#wetlands#erosion control#showy fruits#stream banks#pond margins#specialized bees#cpp#fire medium flammability#NC native#cream flowers#pond garden#native garden#acidic soils tolerant#pollinator plant#naturalized area#nesting sites#larval host plant#food source summer#food source fall#food source herbage#food source nectar#food source pollen#Coastal FACW#Piedmont Mountains FACW#wet soils tolerant#bird friendly#food source soft mast fruit#butterfly friendly#bee friendly#black walnut toxicity tolerant#Audubon#woodland garden#hedge#stormwater garden#wildlife friendly

Attributes:
Genus:

Species:

Family:

Uses (Ethnobotany):

The bark was used by Native Americans as tobacco.

Life Cycle:

Recommended Propagation Strategy:

Stem Cutting

Country Or Region Of Origin:

Eastern Central & Eastern U.S.A

Distribution:

Native: Canada--Ontario and New Foundland: US--AL, CT, DE, DC, FL, IL, IN, IO, KY, ME, MD, MA, MI, MS, MO, NH, NJ, NY, NC, OH, PA, RI, SC, TN, VT, VA, and WV

Fire Risk Rating:

medium flammability

Wildlife Value:

It is the host plant for the spring/summer Azure butterflies. Butterflies nectar at its blooms. Its fruits are eaten by songbirds, ruffed grouse, quail, turkey, chipmunks, black bear, foxes, white-tailed deer, skunks, and squirrels. The foliage is browsed by white-tailed deer. Members of the genus Cornus support the following specialized bees: Andrena (Gonandrena) fragilis, Andrena (Gonandrena) integra, and Andrena (Gonandrena) platyparia.

Play Value:

Attracts Pollinators

Wildlife Cover/Habitat

Wildlife Food Source

Wildlife Larval Host

Wildlife Nesting

Particularly Resistant To (Insects/Diseases/Other Problems):

Deer, Erosion, Wet Soil, Black Walnut. Heat, drought, and soil compaction tolerant.

Dimensions:

Height: 6 ft. 0 in. - 12 ft. 0 in.

Width: 6 ft. 0 in. - 12 ft. 0 in.

Fruit:
Fruit Color:

Fruit Value To Gardener:

Display/Harvest Time:

Fruit Type:

Fruit Length:

Fruit Width:

Fruit Description:

The round drupes are 1/4 inch in diameter. They change from white to blue as they ripen. A cluster of drupes droops from greenish or reddish stalks. Displays from August to September.

Flowers:
Flower Color:

Gold/Yellow

Flower Inflorescence:

Flower Value To Gardener:

Flower Bloom Time:

Flower Petals:

4-5 petals/rays

Flower Size:

Flower Description:

The tiny creamy-white flowers are arranged in flat-topped cymes that are up to 2.5 inches across. The individual flowers are about 3/8 inch wide and have four petals, four sepals, and four long stamens. The flower stalks have silky hairs. The blooms appear from May to June.

Leaves:
Woody Plant Leaf Characteristics:

Leaf Color:

Leaf Feel:

Deciduous Leaf Fall Color:

Insignificant

Leaf Type:

Leaf Arrangement:

Leaf Shape:

Elliptical

Leaf Margin:

Hairs Present:

Leaf Length:

Leaf Width:

Leaf Description:

The leaves are simple, opposite, and elliptical to ovate. The margins are entire. Each leaf measures from 2 to 3.5 inches long and 3/4 to 2 inches wide. The upper surface of the leaves is dark green, and the underside is a paler green. Both surfaces have short stiff hairs. The fall foliage may be green, brown, or purplish-red.

Stem:
Stem Color:

Brown/Copper

Purple/Lavender

Red/Burgundy

Stem Is Aromatic:

Stem Lenticels:

Conspicuous

Stem Surface:

Hairy (pubescent)

Stem Description:

The young twigs may be greenish-purple to dark reddish-purple. They are covered with fine hairs. Older branches are dark purple and eventually develop vertical brownish lenticels or pores. The pith is brown on branches over a year old.

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Cornus amomum is often confused with: Plants that fill a similar niche: Phonetic Spelling KOR-nus ah-MOH-mum Description Silky dogwood is a native large to a medium-sized deciduous shrub in the Cornaceae (dogwood) family. It is native to central and the eastern part of the United States and parts of Canada and is typically found in swamp borders, wetlands, and near streams and ponds.

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Phonetic Spelling

yoo-puh-TOR-ee-um per-foh-lee-AH-tum

This plant has low severity poison characteristics.

Description

Boneset is a large herbaceous, clump-forming rhizomatous perennial herb in the aster (Asteraceae) family that is native to the eastern USA and Canada and as far west as TX, NB and the Dakotas. It can be found in various kinds of wetlands, including marshes, bogs, fens, seeps, edges of rivers, sandy flats and ditches. The plant will grow 4-6 feet tall with a 2-4 foot spread and can form colonies.

The Latin name, Eupatorium, is derived from Eupator, a 1st century BCE king of Pontus, famed for his herbal skills. The species name, perfoliatum, means "through the leaf" since the stem seems to grow through the paired leaves.

Boneset has small white flowers that appear in late summer and fall. The plant grows well in average, medium to wet soils with a consistent water source. It prefers full sun or part shade and tolerates both sandy and clay soils with plenty of organic matter. It will tolerate short-term occasional flooding.

Use this plant in the native pollinator garden and wetland areas such as ponds, streams, marshes or rain and water gardens.

This plant was selected as the 2003 NC Wildflower of the Year, a program managed by the North Carolina Botanical Garden with some financial support from the Garden Club of North Carolina.

Diseases, Insects, and Other Plant Problems: No serious insect or disease problems. Foliage may scorch if soils are allowed to dry out.

See this plant in the following landscape:

Native/ Woodland Garden

Cultivars / Varieties:

Tags:

#poisonous#water garden#fall flowers#white flowers#native perennials#native shrub#NC native#summer flowers#native garden#native wildflower#food source fall#NC Native Pollinator Plant#food source herbage#food source nectar#food source pollen#Coastal FACW#Piedmont Mountains FACW#wet soils tolerant#food source hard mast fruit#butterfly friendly#Audubon#NC Wildflower of the Year#perennial

Cultivars / Varieties:

Tags:

#poisonous#water garden#fall flowers#white flowers#native perennials#native shrub#NC native#summer flowers#native garden#native wildflower#food source fall#NC Native Pollinator Plant#food source herbage#food source nectar#food source pollen#Coastal FACW#Piedmont Mountains FACW#wet soils tolerant#food source hard mast fruit#butterfly friendly#Audubon#NC Wildflower of the Year#perennial

Attributes:
Genus:

Eupatorium

Species:

perfoliatum

Family:

Asteraceae

Uses (Ethnobotany):

Historically used in herbal medicine for the treatment of cases of flu, fevers, colds, and a variety of other maladies.

Life Cycle:

Recommended Propagation Strategy:

Country Or Region Of Origin:

Eastern and central USA and Canada

Distribution:

AL , AR , CT , DC , DE , FL , GA , IA , IL , IN , KS , KY , LA , MA , MD , ME , MI , MN , MO , MS , NC , ND , NE , NH , NJ , NY , OH , OK , PA , RI , SC , SD , TN , TX , VA , VT , WI , WV Canada: MB , NB , NS , ON , PE , QC

Fire Risk Rating:

low flammability

Wildlife Value:

Attracts butterflies and bees. Various moths use this plant as a host. Some birds will eat the seeds

Edibility:

Toxic and bitter

Dimensions:

Height: 4 ft. 0 in. - 6 ft. 0 in.

Width: 3 ft. 0 in. - 4 ft. 0 in.

Fruit:
Display/Harvest Time:

Fruit Type:

Fruit Description:

Achenes have small tufts of hair for wind dispersal. Fruit displays from September to November.

Flowers:
Flower Color:

Flower Inflorescence:

Flower Value To Gardener:

Flower Bloom Time:

Flower Shape:

Flower Size:

Flower Description:

Tiny fragrant white tubular disc florets occur in clusters of 10-20. Numerous small clusters form larger clusters that are 2-8 inches across and occur at the tops of the stems and upper leaf axils. Each floret has 5 spreading petal-like lobes and a long 2-parted, thin style. The bracts at the base of the flowers are light green with some white at the tips. They are in 1-2 layers and are oblong, hairy and gland-dotted. Bloom time is July to Sept.

Leaves:
Leaf Color:

Leaf Feel:

Leaf Type:

Leaf Arrangement:

Leaf Shape:

Lanceolate

Leaf Margin:

Hairs Present:

Leaf Length:

Leaf Width:

Leaf Description:

The yellow-green leaves have a wrinkled look, are finely toothed, and their bases join to surround the stem. Lower leaves are larger, up to 8 inches long and 2 inches wide. Veins are conspicuous and undersides are hairy. Upper leaves are smaller and sessile.

Poisonous to Humans:
Poison Severity:

Poison Symptoms:

It causes vomiting and has a laxative effect laxative in large doses. May cause dermatitis. Potential for liver damage if used long-term. Drying the plant parts helps to decrease toxins.

Poison Toxic Principle:

pyrrolizidine alkaloids

Causes Contact Dermatitis:

Poison Part:

NC State University and N.C. A&T State University work in tandem, along with federal, state and local governments, to form a strategic partnership called N.C. Cooperative Extension, which staffs local offices in all 100 counties and with the Eastern Band of Cherokee Indians.

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N.C. Cooperative Extension prohibits discrimination and harassment on the basis of race, color, national origin, age, sex (including pregnancy), disability, religion, sexual orientation, gender identity, and veteran status.

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Phonetic Spelling yoo-puh-TOR-ee-um per-foh-lee-AH-tum This plant has low severity poison characteristics. Description Boneset is a large herbaceous, clump-forming rhizomatous perennial herb in the aster (Asteraceae) family that is native to the eastern USA and Canada and as far west as TX, NB and the Dakotas.

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https://www.gardenorganic.org.uk/expert-advice/garden-management/weeds/weed-datasheets/redshank#:~:text=It%20occurs%20on%20most%20soils,as%20a%20follower%20of%20man.

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Information on habit, biology, persistence & spread for Redshank.

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https://www.pbs.org/wnet/nature/blog/bumblebee-fact-sheet/#:~:text=Geography%3A,to%20New%20Zealand%20and%20Tasmania.

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As a young girl I remember almond pollination was the time of year that meant our winter was over and it was time to get back to work! Not only was I the co-pilot and expert on-the-road coffee pourer, I was the official load "tie-er downer." My Dad would load the bee hives onto our truck with the boom and once a row was on, I'd come behind him and tie it down. Once the load was on, we would spend hours driving to California's Central Valley to the almond orchards. Once there, we would set the bee hives down in spots designated by the grower.

My job was rolling up and stowing the straps and also what I'm sure was the most comical vision to anyone who might see us (although at 2 in the morning in the middle of an almond orchard there never was anyone) I had to hold the boom and run along beside the truck while Dad moved up a dozen rows. Me running in my big work boots...a gangly teen, trying not to swing around on the end of the boom...pure comedy!

Every year, almond pollination signals the beginning of a busy season for bees and their keepers in California. It's the opening act in the annual crop pollination business of bees and a critical time for the industry. Almond pollination is just the start, and from there, bees move on to pollinate a wide variety of other crops like cherries, melons, avocados, olives, and more. In this article, we'll explore the importance of bee pollination in California's agricultural industry and how it impacts our food supply.

What is Pollination?

Pollination is the transfer of pollen from the male part of a flower (the stamen) to the female part of the same or another flower (the stigma). This transfer is essential for plants to reproduce and produce fruits, vegetables, nuts, and seeds. Pollination can occur through self-pollination, wind pollination, or the action of pollinators like bees.

Why is Pollination Important?

Pollination is important because it allows plants to reproduce and produce fruits, vegetables, nuts, and seeds essential for our food supply. With pollination, we would have the variety of fruits and vegetables we enjoy today. In fact, more than 75% of the world's food crops depend on pollinators like bees.

The Role of Bees in Pollination

Bees are one of the most important pollinators in our food chain. Bees visit flowers to collect nectar and pollen, and in doing so, they transfer pollen from the male to the female part of the flower, allowing fertilization. Bees are especially effective pollinators because they have branched hairs on their bodies that trap and carry pollen from flower to flower.

The Importance of Bee Pollination

Bee pollination is essential for producing many crops, including those grown in California. Bees are responsible for pollinating more than 90 different crops and for more than $15 billion in increased crop value each year. With bees, the yield and quality of these crops would be greatly impacted, leading to a decline in food availability and an increase in food prices.
Almond Pollination

Almond pollination is a crucial time for beekeepers in California, as it's the first major pollination event of the year. Almonds are California's largest agricultural export, and the state produces over 80% of the world's almonds. Beekeepers from across the country bring their hives to California's almond orchards to help pollinate the trees, and it's a significant undertaking.

It's estimated that over 2 million colonies of bees are brought to California for almond pollination each year. Almond Pollination is an important time for the bees because, in many cases, the almond pollen is the first pollen the bees are exposed to after winter. The exposure to this pollen "wakes up" the hive. Once the pollen starts coming in, this tells the Queen spring has arrived and it's time to get to work laying eggs and building the colony size up in preparation for nectar gathering which will come next.
Other Crops Pollinated by Bees
Bees are essential for pollinating many crops besides almonds. Here are some of the other crops that are pollinated by bees in California:

Cherries: Cherries are an important crop in California and require pollination to produce fruit. Bees play a crucial role in pollinating cherry trees; with them, cherries' yield and quality would be greatly impacted. In 2022 there were 12 million pounds of cherries harvested in California...that's a lot of pollinated flowers!

Melons: Melons are a popular summer crop in California and are pollinated by bees. Melons require pollination to produce fruit, and without bees, the yield and quality of melons would be significantly impacted. In 2022, California melons were an 8 million dollar crop!

Avocados: Avocados are a valuable crop in California, and bees are essential for pollinating avocado trees. Avocados require cross-pollination to produce fruit, and bees are the primary pollinators for this crop. Interesting factoid about avocado flowers: the flower opens in the female form during one part of the day, then closes and reopens as a male flower!

Olives: Olives are another crop that is pollinated by bees in California. Bees are responsible for pollinating olive trees, which are grown for oil and table olives. Without bees, the yield and quality of olives would be significantly impacted. One flower equals one olive, that's a lot of bees visiting a lot of flowers!

Apples: Apples are an important crop in California and require pollination to produce fruit. Bees are one of the primary pollinators for apple trees; with them, the yield and quality of apples would be greatly impacted.

Blueberries: Blueberries are a valuable crop in California and require pollination to produce fruit. Bees are one of the primary pollinators for blueberry bushes; with them, the yield and quality of blueberries would be greatly impacted.

These are just a few examples of the many crops pollinated by bees in California. With bees, the yield and quality of these crops would be greatly impacted, leading to a decline in food availability and an increase in food prices. We must take steps to support bee populations to ensure the health and diversity of our ecosystems and our food supply.
The Impact of Bee Decline
Unfortunately, bee populations are declining alarmingly due to habitat loss, pesticide use, climate change, and disease. This decline is concerning because it threatens our food supply and the health of our ecosystems. Without bees, many crops would be unable to reproduce, leading to a decline in food availability and variety. Additionally, the loss of bees would significantly impact the animals that rely on these crops for food and habitat.
What Can We Do to Help Bees?

There are several things we can do to help bees and support their populations:

Plant bee-friendly flowers in our gardens and yards: Bees rely on flowers for food and habitat, and planting various flowers that bloom throughout the season can help support their populations. Some bee-friendly flowers include sunflowers, lavender, cosmos, and zinnias.

Avoid using pesticides and herbicides: Pesticides and herbicides can harm bees and other pollinators. If you must use these chemicals, use them sparingly and follow the instructions carefully to avoid harming bees and other beneficial insects.

Support local beekeepers by purchasing their honey and beeswax products: Beekeepers play a crucial role in managing bee populations and supporting them by purchasing their honey and beeswax products can help ensure their continued success.

Provide nesting habitats for bees, such as bee hotels or nesting boxes: Bees need places to nest and raise their young, and providing nesting habitats like bee hotels or nesting boxes can help support their populations.

Educate others about the importance of bees and pollinators in our food chain: Raising awareness about the importance of bees and other pollinators can help build support for their conservation and ensure the health of our ecosystems and food supply.

By taking these steps, we can help support bee populations and ensure their continued role in pollinating crops and maintaining the health of our ecosystems.

The Importance of Bee Pollination Conclusion

In conclusion, bee pollination is critical to California's agricultural industry and food supply. Beekeepers are essential in pollinating crops like almonds, cherries, melons, avocados, olives, and more. Without bees, our food supply would be severely impacted, and the health of our ecosystems would be at risk. By supporting bees, we can help ensure that they continue to play their vital role in pollination.

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Pollination: The Bee's Integral Link in Our Food Chain

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In this article, we'll explore the importance of bee pollination in California's agricultural industry and how it impacts our food supply. Read on to learn more

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https://www.almanac.com/plant/mint
https://www.thecanadianencyclopedia.ca/en/article/mint#:~:text=Most%20species%20of%20mint%20are,since%20at%20least%20300%20BCE.

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Wild mint ( Mentha arvensis ) This forb is distinctive and reminds many people of refreshing rejuvenation during hot summer days. A crushed leaf releases a sharp mint smell, and cooking with the leaves of wild mint adds a sharp fresh taste to your food.