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Papilio memnon, the great Mormon, is a large butterfly native to southern Asia that belongs to the swallowtail family. It is widely distributed and has thirteen subspecies. The female is polymorphic and with mimetic forms. Its range includes north-eastern India (including Sikkim, Assam and Nagaland), Nepal, Bangladesh, Myanmar, Nicobar Islands, Andaman Islands (stragglers only), western, southern and eastern China (including Hainan), Taiwan southern Japan, Ryukyu Islands, Thailand, Laos, Vietnam, Kampuchea, Malaysia and Indonesia (Sumatra, Mentawai Islands, Nias, Batu, Simeulue, Bangka, Java, Kalimantan and the Lesser Sunda Islands).The butterfly is large with a 120 to 150 millimetres (4.7 to 5.9 in) span. It has four male and many female forms, the females being highly polymorphic and many of them being mimics of unpalatable butterflies. This species has been studied extensively for understanding the genetic basis for polymorphy and Batesian mimicry. As many as twenty-six female forms are reported
কালিম । Common Mormon (Papilio polytes) - Male on a mating dance display with a female (Form: romulus)
A common species of swallowtail butterfly (family: papilionidae) widely distributed across Asia. Seen round the year throughout India from plains up to 2000m. This butterfly is known for the mimicry displayed by the numerous polymorphic forms of its females. These are as follows: cyrus, stichius, romulus.
Host Plant: Ixora coccinea; Rangan (রঙ্গন, Rugmini in Hindi, commonly known as the Jungle Geranium, Flame of the Woods, and Jungle Flame from Rubiaceae family) is an exotic bright red flower, bloom as a flower bunch comprises of lot of small red flowers at the top of branch. Each red flower has four petals and holds four yellow stamen(no filament) between the petals. Flower blooms more or less throughout the year, but best during the rainy season.
Tajpur Sea Beach, Bay of Bengal
Monsoon Images of Bengal, India
Sigma 150-600mm 5-6.3 DG HSM OS Contemporary plus Nikon D7000
They are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. Butterflies have the typical four-stage insect life cycle. Winged adults lay eggs on the food plant on which their larvae, known as caterpillars, will feed. The caterpillars grow, sometimes very rapidly, and when fully developed, pupate in a chrysalis. When metamorphosis is complete, the pupal skin splits, the adult insect climbs out, and after its wings have expanded and dried, it flies off. Some butterflies, especially in the tropics, have several generations in a year, while others have a single generation, and a few in cold locations may take several years to pass through their entire life cycle. Butterflies are often polymorphic, and many species make use of camouflage, mimicry and aposematism to evade their predators. Some, like the monarch and the painted lady, migrate over long distances. Many butterflies are attacked by parasites or parasitoids, including wasps, protozoans, flies, and other invertebrates, or are preyed upon by other organisms. 6446
Innenpark. Bonifatius-Felsen. Felsen am Bonifatiusweg,
Inner Park. Boniface Rock. Rock on the Boniface-Way
Asplenium trichomanes subsp. pachyrachis (Christ) Lovis et Reichst.
Willdenowia 10: 18. 1980.
Asplenium trichomanes sublus. pachyrachis Christ
Farnkr. Schweiz 1 (2): 92. 1900.
Asplenium csikii Kümmerle & András.
Magyar Bot. Lapok 17. 110. 1919 (nomen), 20. 3, fig. 1923
Asplenium pachyrhachis Landolt
Fl. Indicativa 268 2010.
Seestern-Braunschwarz-Streifenfarn, Dickstieliger Braunstieliger Streifenfarn, Dickstieliger Brauner Streifenfarn
Lobed Maidenhair Spleenwort
Anmerkung:
Die Asplenium trichomanes Gruppe, ist ein sehr polymorpher, taxonomisch kritischer Spezieskomplex! Die Evolutionsgeschichte und Beziehungen zwischen den Taxa in dieser Gruppe wurden intensiv untersucht. Allerdings sind morphologische Variation und die Verteilung dieser Taxa unzureichend bekannt, da sie in der lokalen Floren oder Checklisten häufig nicht vorkommen. Die Gründe für die Vernachlässigung der Taxa innerhalb der Asplenium trichomanes Gruppe sind der Mangel an diagnostischen morphologische Merkmalen, das häufige gemeinsame Auftreten an ihren Standorten, sowie die Hybridisierung unter den Taxa. Die Asplenium trichomanes Gruppe umfasst zytologisch und ökologisch unterschiedliche Taxa mit fast weltweiter Verbreitung, die offenbar noch aktiv in der Entwicklung sind (L. Ekrt & M. Štech, 2008).
Annotation:
The Asplenium trichomanes group is a very polymorphic, taxonomically critical species complex! The evolutionary history and relationships between the taxa in this group have been extensively studied. However, morphological variation and distribution of these taxa are poorly understood because they are often absent in local floras or checklists. The reasons for the neglect of the taxa within the Asplenium trichomanes group are the lack of diagnostic morphological features, the frequent common occurrence at their sites, as well as the hybridization among the taxa. The Asplenium trichomanes group includes cytologically and ecologically diverse taxa with almost worldwide distribution, which apparently are still active in development (L. Ekrt & M. Štech, 2008).
Papillons en liberté 2016 / Jardin botanique de Montréal
Distribution : Amériques centrale et du Sud. Envergure des ailes : 7-8 cm. La zone colorée qui irradie vers le bord externe des ailes postérieures peut être rouge vif, orangée, vert d’eau, bleu poudre, ou tellement foncée qu’elle est à peine visible.
* Explore 07/03/2016 *
IMG_4292
It is one of the most common large species of land snail in Europe. size: 18–25 mm
Cepaea nemoralis is highly polymorphic in shell colour and banding. The background colour of the shell varies along a continuum from brown through pink to yellow and sometimes almost white.Additionally the shells can be with or without dark bands. The bands vary in intensity of colour, in width and in number, from zero to five.
This beauty which was temporarily resting on the ground at the Haga Ocean butterfly house in Solna, Sweden is a female common mormon (Papilio polytes f. polytes) - as in the form polytes of the species polytes.
The females of this species are polymorphic and mimic other, unpalatable species. This one makes a fairly good impression of the common rose (Pachliopta aristolochiae), a beautiful species which I haven't photographed.
Part 1 here: www.flickr.com/photos/tinyturtle/53593612199/
Part 2 here: www.flickr.com/photos/tinyturtle/53938375125/
Part 3 here: www.flickr.com/photos/tinyturtle/54340723162/
There is also a colour form which looks just like the males so here is a shot which I am unable to tell if it is a male or female: www.flickr.com/photos/tinyturtle/52907390990/
A destructive polymorphic virus damages a firewall so it can access financial data in an online bank account.
More techno-abstract.
Alternate title:
All uR M0n1E$ r b3L0nG 2 u5!!!1!!!
Enjoy! :)
Sony Alpha 65
Carl Zeiss Vario-Sonnar T* DT 16-80mm F3.5-4.5 ZA
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The Great Mormon (Papilio memnon) is a large butterfly that belongs to the swallowtail family and is found in southern Asia. It is widely distributed and has thirteen subspecies. The female is polymorphic and with mimetic forms.
Tenerife.
Icod, the Butterfly Garden.
Mariposario del Drago.
www.mariposario.com/English/index.html
Great Mormon (Papilio memnon) is a large butterfly with contrasting colors that belongs to the Swallowtail family. A common South-Asian butterfly, it is widely distributed and has thirteen subspecies. The female is polymorphic and with mimetic forms.
Butterflies are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprises the large superfamily Papilionoidea, which contains at least one former group, the skippers (formerly the superfamily "Hesperioidea"), and the most recent analyses suggest it also contains the moth-butterflies (formerly the superfamily "Hedyloidea"). Butterfly fossils date to the Paleocene, about 56 million years ago.
Butterflies have the typical four-stage insect life cycle. Winged adults lay eggs on the food plant on which their larvae, known as caterpillars, will feed. The caterpillars grow, sometimes very rapidly, and when fully developed, pupate in a chrysalis. When metamorphosis is complete, the pupal skin splits, the adult insect climbs out, and after its wings have expanded and dried, it flies off. Some butterflies, especially in the tropics, have several generations in a year, while others have a single generation, and a few in cold locations may take several years to pass through their entire life cycle.
Butterflies are often polymorphic, and many species make use of camouflage, mimicry and aposematism to evade their predators. Some, like the monarch and the painted lady, migrate over long distances. Many butterflies are attacked by parasites or parasitoids, including wasps, protozoans, flies, and other invertebrates, or are preyed upon by other organisms. Some species are pests because in their larval stages they can damage domestic crops or trees; other species are agents of pollination of some plants. Larvae of a few butterflies (e.g., harvesters) eat harmful insects, and a few are predators of ants, while others live as mutualists in association with ants. Culturally, butterflies are a popular motif in the visual and literary arts.
As usual for owls, female screech-owls are usually larger than the males of their species, with owls of both sexes being compact in size and shape.
The Eastern Screech-owl, Megascops asio, is one of the smallest species of owls in North America. All of the birds in this genus are small and agile.
Screech-owls are generally colored in various brownish hues with usually a whitish, patterned underside, which helps to camouflage them against the bark of trees. Some are polymorphic, occurring in a grayish- and a reddish-brown morph.
Went shooting again with www.raymondbarlow.com/
The variable oystercatcher (Haematopus unicolor) is a species of wader in the family Haematopodidae. It is endemic to New Zealand. The Maori name is torea-pango. They are also known as 'red bills'.
"Variable" refers to the frontal plumage, which ranges from pied through mottled to all black. They are polymorphic meaning they have different genetic variants.
Blacker birds are more common in the south. All Stewart Island variable oystercatchers are black. They have pink legs, an orange eye ring and red beaks. They are often seen in pairs on the coast all around New Zealand. During breeding, the pair will defend their territory, sometimes aggressively. Once mated pairs rarely divorce.
After breeding they may be seen within flocks, or on the edges of flocks, of black and white South Island pied oystercatcher (SIPO) which also have vivid orange beaks. After breeding they may even form small flocks of their own. Males are around 678 grams and females slightly larger at around 724 grams.
Variables can be identified as they are slightly larger than the SIPO - SIPO are around 550 grams. Occasionally totally black but if they are pied (black and white) they can be easily confused with SIPO. The variable species has less definition between the black and the white area, as well as a mottled band on the leading edges of the underwing.
Variables also have a smaller white rump patch which is only a band across the base of the tail rather than a wide wedge shape reaching up to the middle of the back as in the SIPO. When mottled they are sometimes called 'smudgies'.
They feed on molluscs, crabs and marine worms. After heavy rain, they sometime go inland in search of earthworms. They can open a shellfish by either hammering a hole in it or getting the bill between the two shells (of a bivalve) and twisting them apart.
They breed in North Island, South Island, Stewart Island, and Chatham Islands. They do not breed inland or beside rivers although the SIPO does.
This image was taken in Akaroa Harbour on the South Island of New Zealand
Ophyris morisii
Endemismo sardo-corso
Sinonimi: Ophyris aranifera, Ophyris exaltata
Specie polimorfa per una frequente ibridazione con una o due specie, si pensa che si sia evoluta in qualche migliaio di anni; queste ibridazioni le rendono più resistenti delle originarie. Alcuni studiosi le farebbero risalire all Ophyris sphegodes.
Bibliografia: La flora della Sardegna.
Ophyris morisii
Sardinian-Corsican endemism
Synonyms: Ophyris aranifera, Ophyris exaltata
Polymorphic species for frequent hybridization with one or two species, it is thought that has evolved in a few thousand years; these hybrids make them more resistant to the original. Some scholars would date to the Ophyris sphegodes.
Bibliography: The flora of Sardinia.
Ophyris morisii
Endemismo sardo-corso
Sinónimos: Ophyris aranifera, Ophyris exaltata
Especies polimórficas para la hibridación frecuente con una o dos especies, se cree que ha evolucionado en unos pocos miles de años; estos híbridos hacen más resistentes a la original. Algunos estudiosos saldrían a las sphegodes Ophyris.
Bibliografía: La flora de Cerdeña.
Tenerife
Icod de los Vinos
Butterfly garden
Papilio memnon, the great Mormon, is a large butterfly native to southern Asia that belongs to the swallowtail family. It is widely distributed and has thirteen subspecies. The female is polymorphic and with mimetic forms.
Papilio memnon, the great Mormon, is a large butterfly native to southern Asia that belongs to the swallowtail family. It is widely distributed and has thirteen subspecies. The female is polymorphic and with mimetic forms.
The butterfly is large with a 120 to 150 millimetres (4.7 to 5.9 in) span. It has four male and many female forms, the females being highly polymorphic and many of them being mimics of unpalatable butterflies. This species has been studied extensively for understanding the genetic basis for polymorphy and Batesian mimicry. As many as twenty-six female forms are reported.
In the Great Mormon butterfly, papilio agenor agenor, the female is polymorphic having several forms which differ in colour and the presence or absence of "tails".
I've noticed only 2 forms in the females which visit our garden, e.g. this one with tails and a tellow abdomen. This may be the "distantianus" form.
কালিম । Common Mormon (Papilio polytes) - Male on a mating dance with a female (Form: romulus)
A common species of swallowtail butterfly (family: papilionidae) widely distributed across Asia. Seen round the year throughout India from plains up to 2000m. This butterfly is known for the mimicry displayed by the numerous polymorphic forms of its females. These are as follows: cyrus, stichius, romulus.
Host Plant: Ixora coccinea; Rangan (রঙ্গন, Rugmini in Hindi, commonly known as the Jungle Geranium, Flame of the Woods, and Jungle Flame from Rubiaceae family) is an exotic bright red flower, bloom as a flower bunch comprises of lot of small red flowers at the top of branch. Each red flower has four petals and holds four yellow stamen(no filament) between the petals. Flower blooms more or less throughout the year, but best during the rainy season.
Tajpur Sea Beach, Bay of Bengal
Monsoon Images of Bengal, India
Sigma 150-600mm 5-6.3 DG HSM OS Contemporary plus Nikon D7000
Remains of barley (Hordeum vulgare) grains found at archaeological sites in the Fertile Crescent indicate that about 10,000 years ago the crop was domesticated there from its wild relative Hordeum spontaneum. The domestication history of barley is revisited based on the assumptions that DNA markers effectively measure genetic distances and that wild populations are genetically different and they have not undergone significant change since domestication. The monophyletic nature of barley domestication is demonstrated based on allelic frequencies at 400 AFLP polymorphic loci studied in 317 wild and 57 cultivated lines. The wild populations from Israel-Jordan are molecularly more similar than are any others to the cultivated gene pool. The results provided support for the hypothesis that the Israel-Jordan area is the region in which barley was brought into culture. Moreover, the diagnostic allele I of the homeobox gene BKn-3, rarely but almost exclusively found in Israel H. spontaneum, is pervasive in western landraces and modern cultivated varieties. In landraces from the Himalayas and India, the BKn-3 allele IIIa prevails, indicating that an allelic substitution has taken place during the migration of barley from the Near East to South Asia. Thus, the Himalayas can be considered a region of domesticated barley diversification.
Innenpark. Bonifatius-Felsen.
Inner Park. Boniface Rock.
Asplenium trichomanes subsp. pachyrachis (Christ) Lovis et Reichst.
Willdenowia 10: 18. 1980.
Asplenium trichomanes sublus. pachyrachis Christ
Farnkr. Schweiz 1 (2): 92. 1900.
Asplenium csikii Kümmerle & András.
Magyar Bot. Lapok 17. 110. 1919 (nomen), 20. 3, fig. 1923
Asplenium pachyrhachis Landolt
Fl. Indicativa 268 2010.
Seestern-Braunschwarz-Streifenfarn, Dickstieliger Braunstieliger Streifenfarn, Dickstieliger Brauner Streifenfarn
Lobed Maidenhair Spleenwort
Anmerkung:
Die Asplenium trichomanes Gruppe, ist ein sehr polymorpher, taxonomisch kritischer Spezieskomplex! Die Evolutionsgeschichte und Beziehungen zwischen den Taxa in dieser Gruppe wurden intensiv untersucht. Allerdings sind morphologische Variation und die Verteilung dieser Taxa unzureichend bekannt, da sie in der lokalen Floren oder Checklisten häufig nicht vorkommen. Die Gründe für die Vernachlässigung der Taxa innerhalb der Asplenium trichomanes Gruppe sind der Mangel an diagnostischen morphologische Merkmalen, das häufige gemeinsame Auftreten an ihren Standorten, sowie die Hybridisierung unter den Taxa. Die Asplenium trichomanes Gruppe umfasst zytologisch und ökologisch unterschiedliche Taxa mit fast weltweiter Verbreitung, die offenbar noch aktiv in der Entwicklung sind (L. Ekrt & M. Štech, 2008).
Annotation:
The Asplenium trichomanes group is a very polymorphic, taxonomically critical species complex! The evolutionary history and relationships between the taxa in this group have been extensively studied. However, morphological variation and distribution of these taxa are poorly understood because they are often absent in local floras or checklists. The reasons for the neglect of the taxa within the Asplenium trichomanes group are the lack of diagnostic morphological features, the frequent common occurrence at their sites, as well as the hybridization among the taxa. The Asplenium trichomanes group includes cytologically and ecologically diverse taxa with almost worldwide distribution, which apparently are still active in development (L. Ekrt & M. Štech, 2008).
Study area. (A) Temporal sequence of sea otter colonization of Glacier Bay National Park and Preserve and Pleasant Island in Icy Straight, Alaska, USA. (B) Sea otter consumption by wolves occurred in areas with the highest estimated sea otter densities (21, 24), but the relative frequency of sea otter in wolf scats was highest in Glacier Bay and Pleasant Island (26, 28).
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Recovery of a marine keystone predator transforms terrestrial predator–prey dynamics
Gretchen H. Roffler orcid.org/0000-0002-8534-3664 gretchen.roffler@alaska.gov, Charlotte E. Eriksson, Jennifer M. Allen, and Taal Levi orcid.org/0000-0003-1853-8311 taal.levi@oregonstate.eduAuthors Info & Affiliations
Edited by Mary Power, University of California Berkeley, CA; received May 25, 2022; accepted November 7, 2022
January 23, 2023
120 (5) e2209037120
doi.org/10.1073/pnas.2209037120
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Metrics
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Vol. 120 | No. 5
Significance
Abstract
Results
Discussion
Materials and Methods
Data, Materials, and Software Availability
Acknowledgments
Supporting Information
References
Significance
While there has been much effort to understand how sea otter recovery transforms nearshore ecosystems, the effects on terrestrial systems have remained uninvestigated. We documented a transboundary food web interaction resulting from the recolonization of sea otters in Icy Straight, Alaska, which provided an ample marine subsidy to island wolves, and subsequently caused extirpation of a terrestrial ungulate. This defies former predictions of predator–prey dynamics which do not account for increasingly abundant and predictable subsidies such that occur on the forefront of the sea otter recovery wave. Although disruption of terrestrial consumer-resource dynamics was previously unforeseen, it hints at both the possibility that these interactions occurred historically and could also become more widespread as sea otters continue to rebound.
Abstract
Sea otters (Enhydra lutris) and wolves (Canis lupus) are two apex predators with strong and cascading effects on ecosystem structure and function. After decades of recovery from near extirpation, their ranges now overlap, allowing sea otters and wolves to interact for the first time in the scientific record. We intensively studied wolves during 2015 to 2021 in an island system colonized by sea otters in the 2000s and by wolves in 2013. After wolf colonization, we quantified shifts in foraging behavior with DNA metabarcoding of 689 wolf scats and stable isotope analyses, both revealing a dietary switch from Sitka black-tailed deer (Odocoileus hemionus), the terrestrial in situ primary prey, to sea otters. Here we show an unexpected result of the reintroduction and restoration of sea otters, which became an abundant marine subsidy for wolves following population recovery. The availability of sea otters allowed wolves to persist and continue to reproduce, subsequently nearly eliminating deer. Genotypes from 390 wolf scats and telemetry data from 13 wolves confirmed island fidelity constituting one of the highest known wolf population densities and upending standardly accepted wolf density predictions based on ungulate abundance. Whereas marine subsidies in other systems are generally derived from lower trophic levels, here an apex nearshore predator became a key prey species and linked nearshore and terrestrial food webs in a recently deglaciated and rapidly changing ecosystem. These results underscore that species restoration may serve as an unanticipated nutrient pathway for recipient ecosystems even resulting in cross-boundary subsidy cascades.
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A widely recognized ecological phenomenon is the connection of ostensibly distinct systems through the transboundary movement of resources (1). Allochthonous subsidies such as marine resources commonly increase productivity of terrestrial systems through the transport of nutrients by physical or biotic agents, or through movement of organisms (1, 2), and may directly or indirectly affect terrestrial food webs and population dynamics (1, 3). The effect of allochthonous subsidies from a donor system depends upon characteristics of the consumer and the focal habitat; more mobile and generalist consumers inhabiting permeable ecosystem boundaries or landscapes with a higher edge to area ratio may receive increased subsidies (1, 4, 5). The magnitude and the timing of the availability of marine subsidies is also influential to predator–prey dynamics. Seasonal availability may result in prey switching, or temporarily satiate predators, easing pressure on resident prey (i.e., apparent mutualism) (6), whereas consistent availability is likely to decouple predator density from local prey availability, resulting in increased predation pressure on resident terrestrial prey (i.e., apparent competition, refs. 3, 7, and 8).
Marine subsidies to large mammalian predators have the potential to be particularly influential given the strong effect apex predators have on their herbivore prey with consequences for vegetation structure and composition. Although wolves (Canis lupus) are considered to be obligate ungulate predators (9) with population densities consistently linked to ungulate density (9, 10), they display a high degree of dietary plasticity and consume a variety of alternative prey (10) including marine resources (11–15). If marine resources are abundant and predictable in space and time, and do not present a risk to obtain, they may allow canid populations to persist despite low abundance of primary prey, which may in effect uncouple their numerical response from ungulate abundance (16–18) leading to apparent competition through increased ungulate predation (19).
Sea otters were once nearly extirpated throughout their North Pacific range but have rapidly recovered in some areas due to reintroduction efforts and legal protection (20, 21). These conservation successes have been hailed not only for recovering an endangered species, but also for restoring a keystone species interaction in nearshore communities (22). Sea otters play an important role as predators of marine invertebrates such that their absence leads to the proliferation of sea urchins, which eliminate kelp forests with concomitant declines in biodiversity (23). Where sea otters recover, they can become extremely abundant. For example, after translocation from the Aleutian Islands in the 1960s, sea otters reached the southern fjord entrance to Glacier Bay National Park in the late 1980s and have continued to expand rapidly (Fig. 1A) before recently approaching local carrying capacity and a population estimate of 8,108 individuals (95% CIs = 6,374, 10,456) (Fig. 2B) in 2018 (24, 25).
Fig. 1.
Study area. (A) Temporal sequence of sea otter colonization of Glacier Bay National Park and Preserve and Pleasant Island in Icy Straight, Alaska, USA. (B) Sea otter consumption by wolves occurred in areas with the highest estimated sea otter densities (21, 24), but the relative frequency of sea otter in wolf scats was highest in Glacier Bay and Pleasant Island (26, 28).
Fig. 2.
Prey population and wolf diet trends. (A) Sitka black-tailed deer abundance and harvest declined on Pleasant Island after wolves became established in 2013. (B) Annual population estimates of sea otters (with 95% credible intervals) in Glacier Bay have increased dramatically since re-establishment. (C) The proportion of sea otters in Pleasant Island wolf diets increased as the proportion of deer decreased, measured by the relative frequency of occurrence of prey identified in scats (D) Isoplot of δ13C and δ15N for Pleasant Island wolf hair indicates increasing relative importance of marine resources in comparison to terrestrial resources coinciding temporally with declines in deer abundance.
We have recently discovered that this profusion of sea otters also serves as a substantial food resource for wolves in the archipelagic landscape of Southeast Alaska and particularly in areas adjacent to Glacier Bay (Fig. 1B) (26, 27) where sea otters are protected and the most abundant marine mammal (25). After decades of recovery, the ranges of wolves and sea otters now overlap allowing these species to interact for the first time in the scientific record. This presents the possibility that the ongoing recolonization of sea otters will, surprisingly, modify species interactions in terrestrial systems and would represent an unusual species interaction in which a keystone marine apex predator also serves as an important bottom-up resource to a terrestrial apex predator.
We report on a serendipitous opportunity to study sea otter-wolf-ungulate interactions focused on Pleasant Island and the adjacent mainland Gustavus Forelands on the periphery of Glacier Bay (Fig. 1A), beginning shortly after wolves became established on Pleasant Island in 2013. From 2015 to 2020, we used two complimentary methods to profile temporal dietary patterns of wolves—DNA metabarcoding of scats and stable isotope analysis of δ13C and δ15N ratios. We evaluated wolf dietary data in combination with population trends of wolves and ungulate prey, human harvest, and winter severity to provide insight on the factors driving predator–prey dynamics. Wolves are highly mobile and typically have a more dispersed home range than their ungulate prey, allowing them to take advantage of patchy resource distribution. To determine whether island wolves were accessing prey available in adjacent mainland systems, we assessed the spatial distribution and movement patterns of wolves by analyzing individual wolf relocation data obtained from GPS-collared wolves and wolf scats genotyped at 38 single nucleotide polymorphic markers (SNPs). In small and isolated populations, such as occur on islands, generalist predators may have large impacts on prey species even leading to local extinctions (5, 27, 28). We found that increasingly abundant marine subsidies decoupled predator–prey (i.e., wolf-ungulate) relationships, as predators switched from ungulates to marine resources. The continued annual reproduction of wolves on a primarily sea otter diet suggests that wolves are not currently energetically limited and that apparent competition between marine and terrestrial prey may prevent the recovery of deer.
Great Mormon: Great Mormon (Papilio memnon) is a large butterfly with contrasting colors that belongs to the Swallowtail family. A common South-Asian butterfly, it is widely distributed and has thirteen subspecies. The female is polymorphic and with mimetic forms. The butterfly is large with 120 to 150mm span.This butterfly is found in North Eastern India (including Sikkim, Assam and Nagaland), Nepal, Bangladesh, Bhutan, Myanmar, Nicobar Islands, Andaman Islands (stragglers only), western, southern and eastern China (including Hainan), Taiwan and southern Japan, Ryukyu Islands. Thailand, Laos, Vietnam, Kampuchea, eastern and peninsular Malaysia, Indonesia ( Sumatra, Mentawai Islands, Nias, Batu, Simeulue, Bangka, Java, Kalimantan and the lesser Sunda Islands (except Timor, Wetar, Babar and Tanimbar).
I left the scene as I had found it, with the falcon on its kill. Before the bird settled back on the ground it circled me several times, wondering perhaps if I was one of those strange southern trees it could land upon.
This species is polymorphic, meaning multiple forms or types appear within the total population. With Gyrfalcon, the different forms, phases or morphs are separated by feather colour into white, grey and dark birds. These three broad categories, however, are further subdivided into a confusing array of gradients. The bird in the photo can probably best be characterized as a dark morph, although I have seen even darker birds.
If you compare this photograph with the next one you can see a considerable difference in the colour tone of the bird. In this image I did not alter the white balance in post-processing - straight out of the camera it is cold, grey and slightly bluish. The white balance in the next photo in the series is warmed up, giving the bird a softer brownish look - and an appearance much closer to how I remember seeing things in the field.
Beyond the bird's colour there is also the issue of how to deal with the background - undifferentiated white sky and white snow-covered ground. To expose on the bird I experimented with moving the exposure compensation - up and up some more, then back down. But in the blinding white light I couldn't really check results on the LCD screen, so I just relied on instinct.
If I was a more systematic processor, the backgrounds in each of the photos would have been adjusted to match one another. If you check the photo stream you can see they don't, even though they were all taken at the same place at about the same time. More to learn, as always.
Another well marked form of the polymorphic Variable Wheatear, marked by its white cap (black in nominate form). The variability of this species has perplexed taxonomists and may involve up to three different species, Photo from Thar Desert, India.
Tenagogerris euphrosyne
Family: Gerridae
Order: Hemiptera
The striders are hemipterans or "true-bugs" - insects that feed by piercing and sucking food through the proboscis.
The striders walk on water, thanks to hydrophobic hairs on the legs. The hairs extend across the whole body too, making the insect able to repel splashes of water. They are so successful in their niche, that the strider family, the Gerridae, are widespread with some 1700 species described, 10% of which are marine.
The Gerridae are polymorphic in that they can have wings in one generation, when there might be a need to relocate to a new water body. The next generation may not have wings however, if the current environment is stable.
DSC00248 copy
And who wouldn't give a squeal of delight to see this for the first time?
I know I did.
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The A. papilionacea group consists of just two species, A. papilionacea itself, commonly known as the Pink Butterfly Orchid and the Fan Lipped Orchid, A. collina. The former is a polymorphic species with a wide distribution and these factors unsurprisingly give rise to a significant range of natural and regional variations. First described by Linnaeus as long ago as 1759, the taxon has been intensely studied ever since, leading to the recognition of many forms and the creation of a list of synonyms far too extensive to detail in these pages (no less than 67 in 2013). A genetic study in 1993, comparing sub-species grandiflora, papilionacea and aegaea (then heroica) from around the Mediterranean, determined that the minimal genetic difference's discovered, were consistent with nothing more than separate geographic populations within a single cohesive gene pool and that being simple morphs, sub-species status could not be justified. These results were not universally accepted and new taxa continue to emerge.
A. papilionacea ssp aegaea was named according to the findings of a paper in 2013 by Lewis and Kreutz. which sought to restore some taxonomic order to a state of confusion in respect of the early flowering form of Pink Butterfly Orchid from the Aegean, formerly known variously as A. papilionacea ssp heroica, O. papilionacea ssp heroica or V. papilionacea var aegaea. Space does not permit even a precis of this interesting and thoroughly researched piece of work, other than to record its conclusion, which was that the basionym heroica was invalid as it is more correctly a synonym for A. laxiflora.
A. papilionacea ssp aegaea is a sturdy, dense flowered plant with a large lip, very reminiscent of grandiflora, albeit less broad shouldered. A distinctive feature of aegaea is the way in which the perianth is normally less closed than that of grandiflora and indeed most other sub-species. It ranges throughout the Aegean basin, across the Peloponnese to the Ionian islands in the west and with a relatively small diaspora in Anatolia to the east. It is an early flowerer which in Crete can be in full bloom by late February.
www.orchidsofbritainandeurope.co.uk/Anacamptis%20papilion...
Cladonia phyllophora Hoffm., syn.: Cladonia alcicornis var. phyllophora (Hoffm.) Malbr., Cladonia cervicornis f. phyllophora (Hoffm.) Dalla Torre & Sarnth., Cladonia degenerans (Flörke) Spreng
Family: Cladoniaceae
EN: Felt cladonia, DE: Beblätterte Becherflechte
Slo.: no name found
Dat.: Sept. 18. 2008
Lat.: 46.32403 Long.: 13.58408
Code: Bot_0297/2008_DSC3510
Habitat: Steep mountain slope, northwest aspect; among large boulders of a recent, large sock slide; in half shade; on sandy, calcareous ground; moderately humid place; protected from direct rain by overhanging rock; average precipitations ~ 3.000 mm/year, average temperature 7-9 deg C, elevations 750 m (2.450 feet), alpine phytogeographical region.
Substratum: sandy soil/raw hummus, among large calcareous boulders.
Place: Bovec basin, Northwest slopes of Mt. Javoršček, 1557 m; toward the end of a dirt forest road, East Julian Alps, Posočje, Slovenia EC.
Comment (relates to Flickr album Cladonia phyllophora): Browsing literature to determine the name of this find I've found only one or two candidates with podetia, which sometimes proliferate in more than two stores from cup margins. Cladonia rappii as well as Cladonia cervikornis/verticilata look similarly from far, but proliferate strictly from the center of the cups. Cladonia ramulosa may look similar too, but rarely (if at all) proliferates in more than two stores and is usually fertile with numerous conspicuous brown apothecia. None of several specimens found in this observation had podetia with apothecia.
The best, although not ideal, fit I've found seems to be Cladonia phyllophora. All sources agree that this taxon is highly polymorphic (google the pictures of it!). The taxon is also very variously interpreted by the authors (Ref. 7.). The description in literature, which seems the closest to this find, is in Brodo, Sharnoff, Sharnoff (2001) (Ref. 2.) mentioning gradually broadening and seemingly soft near the apex podetia having a slightly puffed-up aspect and cup margins richly decorated by small and thick squamules (see Fig. 4.) and brown pycnidia /see Fig.7.). The description in Smith at al (2009) (Ref. 1.) fits reasonably well too, particularly the description of the habit stated as 'often extensive more or less interlocking tiers of proliferating podetia'. However, many sources mention that the surface of the podetia near the base should be areolate with contrasting blackened decorticated and maculated areas (Ref. 1., Ref. 8.) or blackish podetia base (Ref. 7.), which is not the case in this find. Also substratum is usually cited as acid. This find apparently grew on a mixture of sandy soil and raw hummus deposited in gaps among large rock boulders (a few meters across) of a relatively recent large mountain rock slide. It seems possible that it was at least to some extent acid, however, the bedrock and the boulders themselves are no doubt calcareous. I am not sure my determination is correct, but, I am also not aware of a better alternative.
Ref.:
(1) C.W.Smith, et all, The lichens of Great Britain and Ireland,The British Lichen
Society,(2009), p 333.
(2) I.M. Brodo, S.D. Sharnoff, S.Sharnoff, Lichens of North America, Yale Uni. Press (2001), p 265.
(3) V. Wirth, Die Flechten Baden-Württembergs, Teil.1., Ulmer (1995), p 332.
(4) www.researchgate.net/publication/228358096_The_lichen_gen... (accessed May. 31. 2021)
(5) v3.boldsystems.org/index.php/Taxbrowser_Taxonpage?taxid=6... (accessed June 8. 2021)
(6) www.sharnoffphotos.com/lichensB/cladonia_phyllophora.html (accessed June 12. 2021)
(7) www.lichensmaritimes.org/index.php?task=fiche&lichen=... (accessed June 12. 2021)
(8) italic.units.it/index.php?procedure=taxonpage&num=814 (accessed June 14. 2021)
Unit TA-01
Toa:Fire
Safeguard build
Hau – active, stage 8 (polymorphic)
Inferno Sword
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Decided to try to see if the lights in the kitchen could substitute for daylight in a pinch - results are not great but I think I finally have a cheap idea of how I can muzzle my flash for future pictures.
en.wikipedia.org/wiki/Amazonite
"Amazonite, also known as amazonstone,[4] is a green tectosilicate mineral, a variety of the potassium feldspar called microcline.[4][5][6] Its chemical formula is KAlSi3O8,[1][7] which is polymorphic to orthoclase.
Its name is taken from that of the Amazon River, from which green stones were formerly obtained, though it is unknown whether those stones were amazonite.[4] Although it has been used for jewellery for well over three thousand years, as attested by archaeological finds in Middle and New Kingdom Egypt[8] and Mesopotamia, no ancient or medieval authority mentions it. It was first described as a distinct mineral only in the 18th century.[9]"
My daughter is interested in gems, minerals, rocks, and stones. We took a daytrip to Tucson. It was our first time to the shows. There are actually something like 3 dozen shows around Tucson. We went to the Tucson Convention Center for about 3 hours and then to the Kino Sports Complex for about 1.5 hours. We had hoped to get to the 22nd St show and the GIGM Show on W. Starr Pass but did not have time. I wish I had bought my tickets in advance, it would have saved about 30-40 minutes of waiting in line at the Convention Center. I'm glad I brought a Circular Polarizer to cut some of the glare of the glass exhibit cases.
www.visittucson.org/tucson-gem-mineral-fossil-showcase/
"Every year the world-renowned Tucson Gem, Mineral & Fossil Showcase is like a time portal, a trip around the world, and a treasure hunt all rolled into one. Every winter, more than 65,000 guests from around the globe descend upon Tucson, AZ, to buy, sell, trade, and bear witness to rare and enchanting gems, minerals, and fossils at more than 40 gem show locations across the city. If you're planning a winter visit to Tucson, you won't want to miss this three-week-long event.
"Whether you’re looking for a $5 shimmering crystal necklace or a show-stopping $200,000 crystallized rock from an exotic location, the Tucson Gem, Mineral, & Fossil Shows have something for everyone.
www.visittucson.org/blog/post/gems-and-minerals/
TGMS 2024
Tucson Gem Show 2024
Unit LE-01
Toa:Air
Safeguard build
Miru – active, stage 8 (polymorphic)
Cyclone Axe
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Decided to try to see if the lights in the kitchen could substitute for daylight in a pinch - results are not great but I think I finally have a cheap idea of how I can muzzle my flash for future pictures.
Jewel Scarab (Chrysina sp.) - El Oro Province, Ecuador
These colorful beetles were a fun subject to attempt to photograph one drizzly night in the foothills of the Ecuadorian Andes. Stunningly gorgeous beetlese these scarabs would fly in towards lights and on some nights were present in large numbers. Certain species of this genus are polymorphic meaning that their phenotype can be expressed in different ways, so basically there can be different color variants of these beetles. Apparently some varieties of some species are considered quite rare and valuable, and collectors will pay a hefty sum for them. I only saw one individual with what I would call an atypical phenotype and I will post that one eventually. A light green as seen in this beetle was the "wild type" or prevalent color morph in this population atleast. Interestingly all the beetles in this genus seem to be found at higher elevation from 1000-3000 m asl, the ones I encountered were all at about 1000m and were found in a mixed habitat of pasture and fragmented cloud forest.
Unit KO-01
Toa:Ice
Safeguard build
Akaku – active, stage 8 (polymorphic)
Blizzard Sword/Shield
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
Decided to try to see if the lights in the kitchen could substitute for daylight in a pinch - results are not great but I think I finally have a cheap idea of how I can muzzle my flash for future pictures.
One of the most interesting colouration and patterning I've seen on this otherwise very common viper.
Photo from Río Bigal reserve, Ecuador.
Photographed while walking at San Antonio Open Space Preserve, Los Altos, California
Please click on the photo or press the L key to view the larger size
This beautiful Red-tailed Hawk was perched on a horizontal branch, no more than 50 feet from a heavily-used trail that winds up the hillside from the parking area. Many hikers and runners passed this hawk in both directions without noticing the hawk. The hawk itself was constantly moving its head about as it was searching for prey and in this photo was looking up the hill behind it at some movement that had attracted its attention.
Canon 7D Mark II f/5.6 1/500 ISO 400
=======================
From Wikipedia: The red-tailed hawk (Buteo jamaicensis) is a bird of prey that breeds throughout most of North America, from the interior of Alaska and northern Canada to as far south as Panama and the West Indies. It is one of the most common members within the genus of Buteo in North America or worldwide. The red-tailed hawk is one of three species colloquially known in the United States as the "chickenhawk," though it rarely preys on standard-sized chickens. The bird is sometimes also referred to as the red-tail for short, when the meaning is clear in context.
Red-tailed hawks can acclimate to all the biomes within their range, occurring on the edges of non-ideal habitats such as dense forests and sandy deserts. The red-tailed hawk occupies a wide range of habitats and altitudes including deserts, grasslands, coniferous and deciduous forests, agricultural fields and urban areas. Its latitudinal limits fall around the tree line in the Arctic and the species is absent from the high Arctic. It is legally protected in Canada, Mexico and the United States by the Migratory Bird Treaty Act.
The 14 recognized subspecies vary in appearance and range, varying most often in color, and in the west of North America, red-tails are particularly often strongly polymorphic, with individuals ranging from almost white to nearly all black. The subspecies Harlan's hawk (B. j. harlani) is sometimes considered a separate species (B. harlani). The red-tailed hawk is one of the largest members of the genus Buteo, typically weighing from 690 to 1,600 g (1.5 to 3.5 lb) and measuring 45–65 cm (18–26 in) in length, with a wingspan from 110–141 cm (3 ft 7 in–4 ft 8 in). This species displays sexual dimorphism in size, with females averaging about 25% heavier than males.
The diet of red-tailed hawks is highly variable and reflects their status as opportunistic generalist, but in North America, it is most often a predator of small mammals such as rodents. Prey that is terrestrial and diurnal is preferred so types such as ground squirrels are preferential where they naturally occur. Large numbers of birds and reptiles can occur in the diet in several areas and can even be the primary foods. Meanwhile, amphibians, fish and invertebrates can seem rare in the hawk’s regular diet; however, they are not infrequently taken by immature hawks.
Red-tailed hawks may survive on islands absent of native mammals on diets variously including invertebrates such as crabs, or lizards and birds. Like many Buteo, they hunt from a perch most often but can vary their hunting techniques where prey and habitat demand it. Because they are so common and easily trained as capable hunters, the majority of hawks captured for falconry in the United States are red-tails. Falconers are permitted to take only passage hawks (which have left the nest, are on their own, but are less than a year old) so as to not affect the breeding population. Adults, which may be breeding or rearing chicks, may not be taken for falconry purposes and it is illegal to do so. Passage red-tailed hawks are also preferred by falconers because these younger birds have not yet developed the adult behaviors which would make them more difficult to train.
Description:
Red-tailed hawk plumage can be variable, depending on the subspecies and the region. These color variations are morphs, and are not related to molting. The western North American population, B. j. calurus, is the most variable subspecies and has three main color morphs: light, dark, and intermediate or rufous. The dark and intermediate morphs constitute 10–20% of the population in the western United States but seem to constitute only 1-2% of B. j. calurus in western Canada. A whitish underbelly with a dark brown band across the belly, formed by horizontal streaks in feather patterning, is present in most color variations. This feature is variable in eastern hawks and generally absent in some light subspecies (i.e. B. j. fuertesi).
Most adult red-tails have a dark brown nape and upper head which gives them a somewhat hooded appearance, while the throat can variably present a lighter brown “necklace”. Especially in younger birds, the underside may be otherwise covered with dark brown spotting and some adults may too manifest this stippling. The back is usually a slightly darker brown than elsewhere with paler scapular feathers, ranging from tawny to white, forming a variable imperfect “V” on the back. The tail of most adults, which of course gives this species its name, is rufous brick-red above with a variably sized black subterminal band and generally appears light buff-orange from below. In comparison, the typical pale immatures (i.e. less than two years old) typically have a mildly paler headed and tend to show a darker back than adults with more apparent pale wing feather edges above (for descriptions of dark morph juveniles from B. j. calurus, which is also generally apt for description of rare dark morphs of other races, see under that subspecies description). In immature red-tailed hawks of all hues, the tail is a light brown above with numerous small dark brown bars of roughly equal width, but these tend to be much broader on dark morph birds.
Even in young red-tails, the tail may be a somewhat rufous tinge of brown. The bill is relatively short and dark, in the hooked shape characteristic of raptors, and the head can sometimes appear small in size against the thick body frame. The cere, the legs, and the feet of the red-tailed hawk are all yellow, as is the hue of bare parts in many accipitrids of different lineages. Immature birds can be readily identified at close range by their yellowish irises. As the bird attains full maturity over the course of 3–4 years, the iris slowly darkens into a reddish-brown hue, which is the adult eye-color in all races. Seen in flight, adults usually have dark brown along the lower edge of the wings, against a mostly pale wing, which bares light brownish barring. Individually, the underwing coverts can range from all dark to off-whitish (most often more heavily streaked with brown) which contrasts with a distinctive black patagium marking. The wing coloring of adults and immatures is similar but for typical pale morph immatures having somewhat heavier brownish markings.
Though the markings and hue vary across the subspecies, the basic appearance of the red-tailed hawk is relatively consistent. Overall, this species is blocky and broad in shape, often appearing (and being) heavier than other Buteos of similar length. They are the heaviest Buteos on average in eastern North America, albeit scarcely ahead of the larger winged rough-legged buzzard (Buteo lagopus), and second only in size in the west to the ferruginous hawk (Buteo regalis). Red-tailed hawks may be anywhere from the seventh to the ninth heaviest Buteo in the world depending on what figures are used. However, in the northwestern United States, ferruginous hawk females are 35% heavier than female red-tails from the same area. On average, western red-tailed hawks are relatively longer winged and lankier proportioned but are slightly less stocky, compact and heavy than eastern red-tailed hawks in North America. Eastern hawks may also have mildly larger talons and bills than western ones. Based on comparisons of morphology and function amongst all accipitrids, these features imply that western red-tails may need to vary their hunting more frequently to on the wing as the habitat diversifies to more open situations and presumably would hunt more variable and faster prey, whereas the birds of the east, which was historically well-wooded, are more dedicated perch hunters and can take somewhat larger prey but are likely more dedicated mammal hunters. In terms of size variation, red-tailed hawks run almost contrary to Bergmann's rule (i.e. that northern animals should be larger in relation than those closer to the Equator within a species) as one of the northernmost subspecies, B. j. alascensis, is the second smallest race based on linear dimensions and that two of the most southerly occurring races in the United States, B. j. fuertesi and B. j. umbrinus, respectively, are the largest proportioned of all red-tailed hawks. Red-tailed hawks tend have a relatively short but broad tails and thick, chunky wings. Although often described as long winged, the proportional size of the wings is quite small and red-tails have high wing loading for a buteonine hawk. For comparison, two other widespread Buteo hawks in North America were found to weigh: 30 g (1.1 oz) for every square centimeter of wing area in the rough-legged buzzard (Buteo lagopus) and 44 g (1.6 oz) per square cm in the red-shouldered hawk (Buteo lineatus). In contrast, the red-tailed hawk weighed considerably more for their wing area: 199 g (7.0 oz) per square cm.
As is the case with many raptors, the red-tailed hawk displays sexual dimorphism in size, as females are up to 25% larger than males. As is typical in large raptors, frequently reported mean body mass for Red-tailed Hawks are somewhat higher than expansive research reveals. Part of this weight variation is seasonal fluctuations, hawks tending to be heavier in winter than during migration or especially during the trying summer breeding season, and also due to clinal variation. Furthermore, immature hawks are usually lighter in mass than their adult counterparts despite averaging somewhat longer winged and tailed. Male red-tailed hawks may weigh from 690 to 1,300 g (1.52 to 2.87 lb) and females may weigh between 801 and 1,723 g (1.766 and 3.799 lb) (the lowest figure from a migrating female immature from Goshute Mountains, Nevada, the highest from a wintering female in Wisconsin). Some sources claim the largest females can weigh up to 2,000 g (4.4 lb) but whether this is in reference to wild hawks (as opposed to those in captivity or used for falconry) is not clear.[24] The largest known survey of body mass in red-tailed hawks is still credited to Craighead & Craighead (1956), who found 100 males to average 1,028 g (2.266 lb) and 108 females to average 1,244 g (2.743 lb). However, these figures were apparently taken from labels on museum specimens, apparently from natural history collections in Wisconsin and Pennsylvania, without note to the region, age or subspecies of the specimens. However, 16 sources ranging in sample size from the aforementioned 208 specimens to only four hawks in Puerto Rico (with 9 of the 16 studies of migrating red-tails), showed that males weigh a mean of 860.2 g (1.896 lb) and females weigh a mean of 1,036.2 g (2.284 lb), about 15% lighter than prior species-wide published weights. Within the continental United States, average weights of males can range from 840.8 g (1.854 lb) (for migrating males in Chelan County, Washington) to 1,031 g (2.273 lb) (for male hawks found dead in Massachusetts) and females ranged from 1,057.9 g (2.332 lb) (migrants in the Goshutes) to 1,373 g (3.027 lb) (for females diagnosed as B. j. borealis in western Kansas). Size variation in body mass reveals that the red-tailed hawks typically varies only a modest amount and that size differences are geographically inconsistent. Racial variation in average weights of great horned owls (Bubo virginianus) show that mean body mass is nearly twice (the heaviest race is about 36% heavier than the lightest known race on average) as variable as that of the hawk (where the heaviest race is only just over 18% heavier on average than the lightest). Also, great horned owls correspond well at the species level with Bergmann’s rule.
Male red-tailed hawks can reportedly measure 45 to 60 cm (18 to 24 in) in total length, females measuring 48 to 65 cm (19 to 26 in) long. The wingspan typically can range from 105 to 141 cm (3 ft 5 in to 4 ft 8 in), although the largest females may possible span up to 147 cm (4 ft 10 in). In the standard scientific method of measuring wing size, the wing chord is 325.1–444.5 mm (12.80–17.50 in) long. The tail measures 188 to 258.7 mm (7.40 to 10.19 in) in length. The exposed culmen was reported to range from 21.7 to 30.2 mm (0.85 to 1.19 in) and the tarsus averaged 74.7–95.8 mm (2.94–3.77 in) across the races. The middle toe (excluding talon) can range from 38.3 to 53.8 mm (1.51 to 2.12 in), with the hallux-claw (the talon of the rear toe, which has evolved to be the largest in accipitrids) measuring from 24.1 to 33.6 mm (0.95 to 1.32 in) in length.
Identification:
Although they overlap in range with most other American diurnal raptors, identifying most mature red-tailed hawks to species is relatively straightforward, particularly if viewing a typical adult at a reasonable distance. The red-tailed hawk is the only North American hawk with a rufous tail and a blackish patagium marking on the leading edge of its wing (which is obscured only on dark morph adults and Harlan’s hawks by similarly dark colored feathers).
Other larger adult Buteo in North America usually have obvious distinct markings that are absent in red-tails, whether the rufous-brown “beard” of Swainson's hawks (Buteo swainsonii) or the colorful rufous belly and shoulder markings and striking black-and-white mantle of red-shouldered hawks (also the small “windows” seen at the end of their primaries). In perched individuals, even as silhouettes, the shape of large Buteos may be distinctive, such as the wingtips overhanging the tail in several other species, but not in red-tails. North American Buteos range from the dainty, compact builds of much smaller Buteos, such as broad-winged hawk (Buteo platypterus) to the heavyset, neckless look of ferruginous hawks or the rough-legged buzzard which has a compact, smaller appearance than a red-tail in perched birds due to its small bill, short neck and much shorter tarsus, while the opposite effect occurs in flying rough-legs with their much bigger wing area.
In flight, most other large North American Buteo are distinctly longer and slenderer winged than red-tailed hawks, with the much paler ferruginous hawk having peculiarly slender wings in relation to its massive, chunky body. Swainson's hawks are distinctly darker on the wing and ferruginous hawks are much paler winged than typical red-tailed hawks. Pale morph adult ferruginous hawk can show mildly tawny-pink (but never truly rufous) upper tail, and like red-tails tend to have dark markings on underwing-coverts and can have a dark belly band but compared to red-tailed hawks have a distinctly broader head, their remiges are much whiter looking with very small dark primary tips, they lack the red-tail’s diagnostic patagial marks and usually (but not always) also lack the dark subterminal tail-band, and ferruginous have a totally feathered tarsus. With its whitish head, the ferruginous hawk is most similar to Krider's red-tailed hawks, especially in immature plumage, but the larger hawk has broader head and narrower wing shape and the ferruginous immatures are paler underneath and on their legs. Several species share a belly band with the typical red-tailed hawk but they vary from subtle (as in the ferruginous hawk) to solid blackish, the latter in most light-morph rough-legged buzzards. More difficult to identify among adult red-tails are its darkest variations, as most species of Buteo in North America also have dark morphs. Western dark morph red-tails (i.e. calurus) adults, however, retain the typical distinctive brick-red tail which other species lack, which may stand out even more against the otherwise all chocolate brown-black bird. Standard pale juveniles when perched show a whitish patch in the outer half of the upper surface of the wing which other juvenile Buteo lack. The most difficult to identify stages and plumage types are dark morph juveniles, Harlan’s hawk and some Krider’s hawks (the latter mainly with typical ferruginous hawks as aforementioned). Some darker juveniles are similar enough to other Buteo juveniles that it has been stated that they "cannot be identified to species with any confidence under various field conditions." However, field identification techniques have advanced in the last few decades and most experienced hawk-watchers can distinguish even the most vexingly plumaged immature hawks, especially as the wing shapes of each species becomes apparent after seeing many. Harlan’s hawks are most similar to dark morph rough-legged buzzards and dark morph ferruginous hawks. Wing shape is the most reliable identification tool for distinguishing the Harlan’s from these, but also the pale streaking on the breast of Harlan’s, which tends to be conspicuous in most individuals, and is lacking in the other hawks. Also dark morph ferruginous hawks do not have the dark subterminal band of a Harlan’s hawk but do bear a black undertail covert lacking in Harlan’s.
AB2A8653-1_fCAFlkr
The libretto of Tannhäuser combines mythological elements characteristic of German Romantische Oper (Romantic opera) and the medieval setting typical of many French Grand Operas. Wagner brings these two together by constructing a plot involving the 14th-century Minnesingers and the myth of Venus and her subterranean realm of Venusberg. Both the historical and the mythological are united in Tannhäuser's personality; although he is a historical poet composer, little is known about him other than myths that surround him.
Wagner wove a variety of sources into the opera narrative. According to his autobiography, he was inspired by finding the story in "a Volksbuch (popular book) about the Venusberg", which he claimed "fell into his hands", although he admits knowing of the story from the Phantasus of Ludwig Tieck and E. T. A. Hoffmann's story, Der Kampf der Sänger (The Singers' Contest). Tieck's tale, which names the hero "Tannenhäuser", tells of the minnesinger-knight's amorous adventures in the Venusberg, his travels to Rome as a Pilgrim, and his repudiation by the pope. To this Wagner added material from Hoffmann's story, from Serapions-Brüder (1819), describing a song contest at the Wartburg castle,[1] a castle which featured prominently in Thuringian history. Heinrich Heine had provided Wagner with the inspiration for Der fliegende Holländer and Wagner again drew on Heine for Tannhäuser. In Heine's sardonic essay Elementargeister (Elemental spirits), there appears a poem about Tannhäuser and the lure of the grotto of Venus, published in 1837 in the third volume of Der Salon.[1] Other possible sources include Friedrich de la Motte Fouqué's play Der Sängerkrieg auf der Wartburg and Eichendorff's Das Marmorbild (The Marble Statue, 1819).[1][2]
The legend of Tannhäuser, the amorous crusading Franconian knight, and that of the song contest on the Wartburg (which did not involve Tannhäuser, but the semi-mythical minnesinger Heinrich von Ofterdingen), came from quite separate traditions. Ludwig Bechstein wove together the two legends in the first volume of his collection of Thuringian legends, Der Sagenschatz und die Sagenkreise des Thüringerlandes (A treasury of the tales of Thuringian legends and legend cycles, 1835), which was probably the Volksbuch to which Wagner refers to in his autobiography.[3][1] Wagner also knew of the work of another contemporary, Christian Theodor Ludwig Lucas, whose Über den Krieg von Wartburg of 1838 also conflated the two legends.[4][5] This confusion (which explains why Tannhäuser is referred to as 'Heinrich' in the opera) does not fit with the historical timeline of the events in the opera, since the Singers' Contest involving von Ofterdingen is said to have taken place around 1207, while Tannhäuser's poetry appeared much later (1245–1265). The sources used by Wagner therefore reflected a nineteenth century romantic view of the medieval period, with concerns about artistic freedom and the constraints of organised religion typical of the period of Romanticism.[6]
During Wagner's first stay in Paris (1839–1842) he read a paper by Ludwig Lucas on the Sängerkrieg which sparked his imagination, and encouraged him to return to Germany, which he reached on 7 April 1842.[7] Having crossed the Rhine, the Wagners drove towards Thuringia, and saw the early rays of sun striking the Wartburg; Wagner immediately began to sketch the scenery that would become the stage sets.[8] Wagner wrote the prose draft of Tannhäuser between June and July 1842 and the libretto in April 1843.[9]
en.wikipedia.org/wiki/Tannhäuser_(opera)
Neuschwanstein Castle (German: Schloss Neuschwanstein, pronounced [ˈʃlɔs nɔʏˈʃvaːnʃtaɪn], Southern Bavarian: Schloss Neischwanstoa) is a 19th-century Romanesque Revival palace on a rugged hill above the village of Hohenschwangau near Füssen in southwest Bavaria, Germany. The palace was commissioned by King Ludwig II of Bavaria as a retreat and in honour of Richard Wagner. Ludwig paid for the palace out of his personal fortune and by means of extensive borrowing, rather than Bavarian public funds.
The castle was intended as a home for the King, until he died in 1886. It was open to the public shortly after his death.[1] Since then more than 61 million people have visited Neuschwanstein Castle.[2] More than 1.3 million people visit annually, with as many as 6,000 per day in the summer.[3]
Contents
1Location
2History
2.1Inspiration and design
2.2Construction
2.3Funding
2.4Simplified completion
2.5World War II
3Architecture
3.1Exterior
3.2Interior
4Tourism
5In culture, art, and science
5.1World Heritage candidature
6Panoramas
7Notes
8Citations
9General sources
10External links
Location[edit]
A northward view of Neuschwanstein Castle from Mount Säuling (2,047 m or 6,716 ft) on the border between Bavaria and Tyrol: Schwangau between large Forggensee reservoir (1952) and Hohenschwangau and Neuschwanstein palaces
The municipality of Schwangau lies at an elevation of 800 m (2,620 ft) at the southwest border of the German state of Bavaria. Its surroundings are characterised by the transition between the Alpine foothills in the south (toward the nearby Austrian border) and a hilly landscape in the north that appears flat by comparison.
In the Middle Ages, three castles overlooked the villages. One was called Schwanstein Castle.[nb 1] In 1832, Ludwig's father King Maximilian II of Bavaria bought its ruins to replace them with the comfortable neo-Gothic palace known as Hohenschwangau Castle. Finished in 1837, the palace became his family's summer residence, and his elder son Ludwig (born 1845) spent a large part of his childhood here.[4]
Vorderhohenschwangau Castle and Hinterhohenschwangau Castle[nb 2] sat on a rugged hill overlooking Schwanstein Castle, two nearby lakes (Alpsee and Schwansee), and the village. Separated by only a moat, they jointly consisted of a hall, a keep, and a fortified tower house.[5] In the nineteenth century only ruins remained of the twin medieval castles, but those of Hinterhohenschwangau served as a lookout place known as Sylphenturm.[6]
The ruins above the family palace were known to the crown prince from his excursions. He first sketched one of them in his diary in 1859.[7] When the young king came to power in 1864, the construction of a new palace in place of the two ruined castles became the first in his series of palace building projects.[8] Ludwig called the new palace New Hohenschwangau Castle; only after his death was it renamed Neuschwanstein.[9] The confusing result is that Hohenschwangau and Schwanstein have effectively swapped names: Hohenschwangau Castle replaced the ruins of Schwanstein Castle, and Neuschwanstein Castle replaced the ruins of the two Hohenschwangau Castles.
History[edit]
Inspiration and design[edit]
Neuschwanstein embodies both the contemporaneous architectural fashion known as castle romanticism (German: Burgenromantik), and King Ludwig II's enthusiasm for the operas of Richard Wagner.
In the 19th century, many castles were constructed or reconstructed, often with significant changes to make them more picturesque. Palace-building projects similar to Neuschwanstein had been undertaken earlier in several of the German states and included Hohenschwangau Castle, Lichtenstein Castle, Hohenzollern Castle, and numerous buildings on the River Rhine such as Stolzenfels Castle.[10] The inspiration for the construction of Neuschwanstein came from two journeys in 1867—one in May to the reconstructed Wartburg near Eisenach,[11] another in July to the Château de Pierrefonds, which Eugène Viollet-le-Duc was transforming from a ruined castle into a historistic palace.[12][nb 3]
Neuschwanstein project drawing (Christian Jank 1869)
The King saw both buildings as representatives of a romantic interpretation of the Middle Ages, as well as the musical mythology of his friend Wagner, whose operas Tannhäuser and Lohengrin had made a lasting impression on him.[13]
In February 1868, Ludwig's grandfather King Ludwig I died, freeing the considerable sums that were previously spent on the abdicated King's appanage.[8][nb 4] This allowed Ludwig II to start the architectural project of building a private refuge in the familiar landscape far from the capital Munich, so that he could live out his idea of the Middle Ages.
It is my intention to rebuild the old castle ruin of Hohenschwangau near the Pöllat Gorge in the authentic style of the old German knights' castles, and I must confess to you that I am looking forward very much to living there one day [...]; you know the revered guest I would like to accommodate there; the location is one of the most beautiful to be found, holy and unapproachable, a worthy temple for the divine friend who has brought salvation and true blessing to the world. It will also remind you of "Tannhäuser" (Singers' Hall with a view of the castle in the background), "Lohengrin'" (castle courtyard, open corridor, path to the chapel) ...
— Ludwig II, Letter to Richard Wagner, May 1868[14]
The building design was drafted by the stage designer Christian Jank and realised by the architect Eduard Riedel.[15] For technical reasons, the ruined castles could not be integrated into the plan. Initial ideas for the palace drew stylistically on Nuremberg Castle and envisaged a simple building in place of the old Vorderhohenschwangau Castle, but they were rejected and replaced by increasingly extensive drafts, culminating in a bigger palace modelled on the Wartburg.[16] The king insisted on a detailed plan and on personal approval of each and every draft.[17] Ludwig's control went so far that the palace has been regarded as his own creation, rather than that of the architects involved.[18]
Whereas contemporary architecture critics derided Neuschwanstein, one of the last big palace building projects of the nineteenth century, as kitsch, Neuschwanstein and Ludwig II's other buildings are now counted among the major works of European historicism.[19][20] For financial reasons, a project similar to Neuschwanstein – Falkenstein Castle – never left the planning stages.[21]
The palace can be regarded as typical for nineteenth-century architecture. The shapes of Romanesque (simple geometric figures such as cuboids and semicircular arches), Gothic (upward-pointing lines, slim towers, delicate embellishments) and Byzantine architecture and art (the Throne Hall décor) were mingled in an eclectic fashion and supplemented with 19th-century technical achievements. The Patrona Bavariae and Saint George on the court face of the Palas (main building) are depicted in the local Lüftlmalerei style, a fresco technique typical for Allgäu farmers' houses, while the unimplemented drafts for the Knights' House gallery foreshadow elements of Art Nouveau.[22] Characteristic of Neuschwanstein's design are theatre themes: Christian Jank drew on coulisse drafts from his time as a scenic painter.[23]
The basic style was originally planned to be neo-Gothic but the palace was primarily built in Romanesque style in the end. The operatic themes moved gradually from Tannhäuser and Lohengrin to Parsifal.[24]
Construction[edit]
Neuschwanstein under construction: Bower still missing, Rectangular Tower under construction (photograph c. 1882–85)
Neuschwanstein under construction: upper courtyard (photograph c. 1886)
In 1868, the ruins of the medieval twin castles were completely demolished; the remains of the old keep were blown up.[25] The foundation stone for the palace was laid on 5 September 1869; in 1872 its cellar was completed and in 1876, everything up to the first floor, the gatehouse being finished first. At the end of 1882 it was completed and fully furnished, allowing Ludwig to take provisional lodgings there and observe the ongoing construction work.[24] In 1874, management of the civil works passed from Eduard Riedel to Georg von Dollmann.[26] The topping out ceremony for the Palas was in 1880, and in 1884, the King was able to move in to the new building. In the same year, the direction of the project passed to Julius Hofmann, after Dollmann had fallen from the King's favour.
The palace was erected as a conventional brick construction and later encased in various types of rock. The white limestone used for the fronts came from a nearby quarry.[27]
The sandstone bricks for the portals and bay windows came from Schlaitdorf in Württemberg. Marble from Untersberg near Salzburg was used for the windows, the arch ribs, the columns and the capitals. The Throne Hall was a later addition to the plans and required a steel framework.
The transport of building materials was facilitated by scaffolding and a steam crane that lifted the material to the construction site. Another crane was used at the construction site. The recently founded Dampfkessel-Revisionsverein (Steam Boiler Inspection Association) regularly inspected both boilers.
For about two decades the construction site was the principal employer in the region.[28] In 1880, about 200 craftsmen were occupied at the site,[29] not counting suppliers and other persons indirectly involved in the construction. At times when the King insisted on particularly close deadlines and urgent changes, reportedly up to 300 workers per day were active, sometimes working at night by the light of oil lamps. Statistics from the years 1879/1880 support an immense amount of building materials: 465 tonnes (513 short tons) of Salzburg marble, 1,550 t (1,710 short tons) of sandstone, 400,000 bricks and 2,050 cubic metres (2,680 cu yd) of wood for the scaffolding.
In 1870, a society was founded for insuring the workers, for a low monthly fee, augmented by the King. The heirs of construction casualties (30 cases are mentioned in the statistics) received a small pension.
In 1884, the King was able to move into the (still unfinished) Palas,[30] and in 1885, he invited his mother Marie to Neuschwanstein on the occasion of her 60th birthday.[nb 5] By 1886, the external structure of the Palas (hall) was mostly finished.[30] In the same year, Ludwig had the first, wooden Marienbrücke over the Pöllat Gorge replaced by a steel construction.
Despite its size, Neuschwanstein did not have space for the royal court, but contained only the King's private lodging and servants' rooms. The court buildings served decorative, rather than residential purposes:[9] The palace was intended to serve King Ludwig II as a kind of inhabitable theatrical setting.[30] As a temple of friendship it was also dedicated to the life and work of Richard Wagner, who died in 1883 before he had set foot in the building.[31] In the end, Ludwig II lived in the palace for a total of only 172 days.[32]
Funding[edit]
Neuschwanstein in 1886
The King's wishes and demands expanded during the construction of Neuschwanstein, and so did the expenses. Drafts and estimated costs were revised repeatedly.[33] Initially a modest study was planned instead of the great throne hall, and projected guest rooms were struck from the drafts to make place for a Moorish Hall, which could not be realised due to lack of resources. Completion was originally projected for 1872, but deferred repeatedly.[33]
Neuschwanstein, the symbolic medieval knight's castle, was not King Ludwig II's only huge construction project. It was followed by the rococo style Lustschloss of Linderhof Palace and the baroque palace of Herrenchiemsee, a monument to the era of absolutism.[8] Linderhof, the smallest of the projects, was finished in 1886, and the other two remain incomplete. All three projects together drained his resources. The King paid for his construction projects by private means and from his civil list income. Contrary to frequent claims, the Bavarian treasury was not directly burdened by his buildings.[30][34] From 1871, Ludwig had an additional secret income in return for a political favour given to Otto von Bismarck.[nb 6]
The construction costs of Neuschwanstein in the King's lifetime amounted to 6.2 million marks (equivalent to 40 million 2009 €),[35] almost twice the initial cost estimate of 3.2 million marks.[34] As his private means were insufficient for his increasingly escalating construction projects, the King continuously opened new lines of credit.[36] In 1876, a court counselor was replaced after pointing out the danger of insolvency.[37] By 1883 he already owed 7 million marks,[38] and in spring 1884 and August 1885 debt conversions of 7.5 million marks and 6.5 million marks, respectively, became necessary.[36]
Even after his debts had reached 14 million marks, King Ludwig II insisted on continuation of his architectural projects; he threatened suicide if his creditors seized his palaces.[37] In early 1886, Ludwig asked his cabinet for a credit of 6 million marks, which was denied. In April, he followed Bismarck's advice to apply for the money to his parliament. In June the Bavarian government decided to depose the King, who was living at Neuschwanstein at the time. On 9 June he was incapacitated, and on 10 June he had the deposition commission arrested in the gatehouse.[39] In expectation of the commission, he alerted the gendarmerie and fire brigades of surrounding places for his protection.[36] A second commission headed by Bernhard von Gudden arrived on the next day, and the King was forced to leave the palace that night. Ludwig was put under the supervision of von Gudden. On 13 June, both died under mysterious circumstances in the shallow shore water of Lake Starnberg near Berg Castle.
Simplified completion[edit]
Neuschwanstein front façade and surroundings (photochrom print, c. 1900)
A 1901 postcard of Berg Castle
At the time of King Ludwig's death the palace was far from complete. He slept only 11 nights in the castle. The external structures of the Gatehouse and the Palas were mostly finished but the Rectangular Tower was still scaffolded. Work on the Bower had not started, but was completed in a simplified form by 1892 without the planned figures of the female saints. The Knights' House was also simplified. In King Ludwig's plans the columns in the Knights' House gallery were held as tree trunks and the capitals as the corresponding crowns. Only the foundations existed for the core piece of the palace complex: a keep of 90 metres (300 ft) height planned in the upper courtyard, resting on a three-nave chapel. This was not realised,[17] and a connection wing between the Gatehouse and the Bower saw the same fate.[40] Plans for a castle garden with terraces and a fountain west of the Palas were also abandoned after the King's death.
The interior of the royal living space in the palace was mostly completed in 1886; the lobbies and corridors were painted in a simpler style by 1888.[41] The Moorish Hall desired by the King (and planned below the Throne Hall) was not realised any more than the so-called Knights' Bath, which, modelled after the Knights' Bath in the Wartburg, was intended to render homage to the knights' cult as a medieval baptism bath. A Bride Chamber in the Bower (after a location in Lohengrin),[23] guest rooms in the first and second floor of the Palas and a great banquet hall were further abandoned projects.[33] In fact, a complete development of Neuschwanstein had never even been planned, and at the time of the King's death there was not a utilisation concept for numerous rooms.[29]
Neuschwanstein was still incomplete when Ludwig II died in 1886. The King never intended to make the palace accessible to the public.[30] No more than six weeks after the King's death, however, the Prince-Regent Luitpold ordered the palace opened to paying visitors. The administrators of King Ludwig's estate managed to balance the construction debts by 1899.[42] From then until World War I, Neuschwanstein was a stable and lucrative source of revenue for the House of Wittelsbach, indeed King Ludwig's castles were probably the single largest income source earned by the Bavarian royal family in the last years prior to 1914. To guarantee a smooth course of visits, some rooms and the court buildings were finished first. Initially the visitors were allowed to move freely in the palace, causing the furniture to wear quickly.
When Bavaria became a republic in 1918, the government socialised the civil list. The resulting dispute with the House of Wittelsbach led to a split in 1923: King Ludwig's palaces including Neuschwanstein fell to the state and are now managed by the Bavarian Palace Department, a division of the Bavarian finance ministry. Nearby Hohenschwangau Castle fell to the Wittelsbacher Ausgleichsfonds, whose revenues go to the House of Wittelsbach.[43] The visitor numbers continued to rise, reaching 200,000 in 1939.[43]
World War II[edit]
Due to its secluded location, the palace survived the destruction of two World Wars. Until 1944, it served as a depot for Nazi plunder that was taken from France by the Reichsleiter Rosenberg Institute for the Occupied Territories (Einsatzstab Reichsleiter Rosenberg für die besetzten Gebiete), a suborganisation of the Nazi Party.[44] The castle was used to catalogue the works of arts. (After World War II 39 photo albums were found in the palace documenting the scale of the art seizures. The albums are now stored in the United States National Archives.[45])
In April 1945, the SS considered blowing up the palace to prevent the building itself and the artwork it contained from falling to the enemy.[46] The plan was not realised by the SS-Gruppenführer who had been assigned the task, however, and at the end of the war the palace was surrendered undamaged to representatives of the Allied forces.[46] Thereafter the Bavarian archives used some of the rooms as a provisional store for salvaged archivalia, as the premises in Munich had been bombed.[47]
Architecture[edit]
The effect of the Neuschwanstein ensemble is highly stylistic, both externally and internally. The king's influence is apparent throughout, and he took a keen personal interest in the design and decoration. An example can be seen in his comments, or commands, regarding a mural depicting Lohengrin in the Palas; "His Majesty wishes that ... the ship be placed further from the shore, that Lohengrin's neck be less tilted, that the chain from the ship to the swan be of gold and not of roses, and finally that the style of the castle shall be kept medieval."[48]
The suite of rooms within the Palas contains the Throne Room, King Ludwig's suite, the Singers' Hall, and the Grotto. The interior and especially the throne room Byzantine-Arab construction resumes to the chapels and churches of the royal Sicilian Norman-Swabian period in Palermo related to the Kings of Germany House of Hohenstaufen. Throughout, the design pays homage to the German legends of Lohengrin, the Swan Knight. Hohenschwangau, where King Ludwig spent much of his youth, had decorations of these sagas. These themes were taken up in the operas of Richard Wagner. Many rooms bear a border depicting the various operas written by Wagner, including a theatre permanently featuring the set of one such play. Many of the interior rooms remain undecorated, with only 14 rooms finished before Ludwig's death. With the palace under construction at the King's death, one of the major features of the palace remained unbuilt. A massive keep, which would have formed the highest point and central focus of the ensemble, was planned for the middle of the upper courtyard but was never built, at the decision of the King's family. The foundation for the keep is visible in the upper courtyard.[49]
Neuschwanstein Castle consists of several individual structures which were erected over a length of 150 metres on the top of a cliff ridge. The elongate building is furnished with numerous towers, ornamental turrets, gables, balconies, pinnacles and sculptures. Following Romanesque style, most window openings are fashioned as bi- and triforia. Before the backdrop of the Tegelberg and the Pöllat Gorge in the south and the Alpine foothills with their lakes in the north, the ensemble of individual buildings provides varying picturesque views of the palace from all directions. It was designed as the romantic ideal of a knight's castle. Unlike "real" castles, whose building stock is in most cases the result of centuries of building activity, Neuschwanstein was planned from the inception as an intentionally asymmetric building, and erected in consecutive stages.[33] Typical attributes of a castle were included, but real fortifications – the most important feature of a medieval aristocratic estate – were dispensed with.
Exterior[edit]
Palace roof
Overview of palace complex; position of the planned chapel marked in yellow
View from location of unrealised chapel along upper courtyard level: Bower (left), palace front, and Knights' House (right)
The palace complex is entered through the symmetrical Gatehouse flanked by two stair towers. The eastward-pointing gate building is the only structure of the palace whose wall area is fashioned in high-contrast colours; the exterior walls are cased with red bricks, the court fronts with yellow limestone. The roof cornice is surrounded by pinnacles. The upper floor of the Gatehouse is surmounted by a crow-stepped gable and held King Ludwig II's first lodging at Neuschwanstein, from which he occasionally observed the building work before the hall was completed. The ground floors of the Gatehouse were intended to accommodate the stables.
The passage through the Gatehouse, crowned with the royal Bavarian coat of arms, leads directly into the courtyard. The courtyard has two levels, the lower one being defined to the east by the Gatehouse and to the north by the foundations of the so-called Rectangular Tower and by the gallery building. The southern end of the courtyard is open, imparting a view of the surrounding mountain scenery. At its western end, the courtyard is delimited by a bricked embankment, whose polygonally protracting bulge marks the choir of the originally projected chapel; this three-nave church, never built, was intended to form the base of a 90-metre (295-ft) keep, the planned centrepiece of the architectural ensemble. A flight of steps at the side gives access to the upper level.
Saint George
Gatehouse
Today, the foundation plan of the chapel-keep is marked out in the upper-courtyard pavement. The most striking structure of the upper court level is the so-called Rectangular Tower (45 metres or 148 feet). Like most of the court buildings, it mostly serves a decorative purpose as part of the ensemble. Its viewing platform provides a vast view over the Alpine foothills to the north. The northern end of the upper courtyard is defined by the so-called Knights' House. The three-storey building is connected to the Rectangular Tower and the Gatehouse by means of a continuous gallery fashioned with a blind arcade. From the point of view of castle romanticism the Knights' House was the abode of a stronghold's menfolk; at Neuschwanstein, estate and service rooms were envisioned here. The Bower, which complements the Knights' House as the "ladies' house" but was never used as such, defines the south side of the courtyard. Both structures together form the motif of the Antwerp Castle featuring in the first act of Lohengrin. Embedded in the pavement is the floor plan of the planned palace chapel.
The western end of the courtyard is delimited by the Palas (hall). It constitutes the real main and residential building of the castle and contains the King's stateroom and the servants' rooms. The Palas is a colossal five-story structure in the shape of two huge cuboids that are connected in a flat angle and covered by two adjacent high gable roofs. The building's shape follows the course of the ridge. In its angles there are two stair towers, the northern one surmounting the palace roof by several storeys with its height of 65 metres (213 ft). With their polymorphic roofs, both towers are reminiscent of the Château de Pierrefonds. The western Palas front supports a two-storey balcony with view on the Alpsee, while northwards a low chair tower and the conservatory protract from the main structure. The entire Palas is spangled with numerous decorative chimneys and ornamental turrets, the court front with colourful frescos. The court-side gable is crowned with a copper lion, the western (outward) gable with the likeness of a knight.
Interior[edit]
Floor plan of third floor, position of fourth-floor Hall of the Singers marked in red
Corridor
Throne Hall detail
Had it been completed, the palace would have had more than 200 interior rooms, including premises for guests and servants, as well as for service and logistics. Ultimately, no more than about 15 rooms and halls were finished.[50] In its lower stories the Palas accommodates administrative and servants' rooms and the rooms of today's palace administration. The King's staterooms are situated in the upper stories: The anterior structure accommodates the lodgings in the third floor, above them the Hall of the Singers. The upper floors of the west-facing posterior structure are filled almost completely by the Throne Hall. The total floor space of all floors amounts to nearly 6,000 square metres (65,000 sq ft).[50]
Neuschwanstein houses numerous significant interior rooms of German historicism. The palace was fitted with several of the latest technical innovations of the late 19th century.[22][51] Among other things it had a battery-powered bell system for the servants and telephone lines. The kitchen equipment included a Rumford oven that turned the skewer with its heat and so automatically adjusted the turning speed. The hot air was used for a calorifère central heating system.[52] Further novelties for the era were running warm water and toilets with automatic flushing.
The largest room of the palace by area is the Hall of the Singers, followed by the Throne Hall. The 27-by-10-metre (89 by 33 ft)[53] Hall of the Singers is located in the eastern, court-side wing of the Palas, in the fourth floor above the King's lodgings. It is designed as an amalgamation of two rooms of the Wartburg: The Hall of the Singers and the Ballroom. It was one of the King's favourite projects for his palace.[54] The rectangular room was decorated with themes from Lohengrin and Parzival. Its longer side is terminated by a gallery that is crowned by a tribune, modelled after the Wartburg. The eastern narrow side is terminated by a stage that is structured by arcades and known as the Sängerlaube. The Hall of the Singers was never designed for court festivities of the reclusive King.[citation needed] Rather, like the Throne Hall it served as a walkable monument in which the culture of knights and courtly love of the Middle Ages was represented. The first performance in this hall took place in 1933: A concert commemorating the 50th anniversary of Richard Wagner's death.[34]
The Throne Hall, 20 by 12 metres (66 by 39 ft),[55] is situated in the west wing of the Palas. With its height of 13 metres (43 ft)[55] it occupies the third and fourth floors. Julius Hofmann modelled it after the Allerheiligen-Hofkirche in the Munich Residenz. On three sides it is surrounded by colorful arcades, ending in an apse that was intended to hold King Ludwig's throne – which was never completed. The throne dais is surrounded by paintings of Jesus, the Twelve Apostles and six canonised kings. The mural paintings were created by Wilhelm Hauschild. The floor mosaic was completed after the king's death. The chandelier is fashioned after a Byzantine crown. The Throne Hall makes a sacral impression. Following the king's wish, it amalgamated the Grail Hall from Parzival with a symbol of the divine right of kings,[19] an incorporation of unrestricted sovereign power, which King Ludwig as the head of a constitutional monarchy no longer held. The union of the sacral and regal is emphasised by the portraits in the apse of six canonised Kings: Saint Louis of France, Saint Stephen of Hungary, Saint Edward the Confessor of England, Saint Wenceslaus of Bohemia, Saint Olaf of Norway and Saint Henry, Holy Roman Emperor.
Palace rooms (late 19th century Photochrom prints)
Hall of the Singers
Throne Hall
Drawing room
Study room
Dining room
Bedroom
Apart from the large ceremonial rooms several smaller rooms were created for use by King Ludwig II.[41] The royal lodging is on the third floor of the palace in the east wing of the Palas. It consists of eight rooms with living space and several smaller rooms. In spite of the gaudy décor, the living space with its moderate room size and its sofas and suites makes a relatively modern impression on today's visitors. King Ludwig II did not attach importance to representative requirements of former times, in which the life of a monarch was mostly public. The interior decoration with mural paintings, tapestry, furniture and other handicraft generally refers to the King's favourite themes: the grail legend, the works of Wolfram von Eschenbach, and their interpretation by Richard Wagner.
Grotto
The eastward drawing room is adorned with themes from the Lohengrin legend. The furniture – sofa, table, armchairs and seats in a northward alcove – is comfortable and homelike. Next to the drawing room is a little artificial grotto that forms the passage to the study. The unusual room, originally equipped with an artificial waterfall and a so-called rainbow machine, is connected to a little conservatory. Depicting the Hörselberg grotto, it relates to Wagner's Tannhäuser, as does the décor of the adjacent study. In the park of Linderhof Palace the King had installed a similar grotto of greater dimensions. Opposite the study follows the dining room, adorned with themes of courtly love. Since the kitchen in Neuschwanstein is situated three stories below the dining room, it was impossible to install a wishing table (dining table disappearing by means of a mechanism) as at Linderhof Palace and Herrenchiemsee. Instead, the dining room was connected with the kitchen by means of a service lift.
Kitchen
The bedroom adjacent to the dining room and the subsequent house chapel are the only rooms of the palace that remain in neo-Gothic style. The King's bedroom is dominated by a huge bed adorned with carvings. Fourteen carvers worked more than four years on the bed canopy with its numerous pinnacles and on the oaken panellings.[56] It was in this room that Ludwig was arrested in the night from 11 to 12 June 1886. The adjacent little house chapel is consecrated to Saint Louis, after whom the owner was named.
The servants' rooms in the basement of the Palas are quite scantily equipped with massive oak furniture. Besides one table and one cabinet there are two beds of 1.80 metres (5 ft 11 in) length each. Opaque glass windows separated the rooms from the corridor that connects the exterior stairs with the main stairs, so that the King could enter and leave unseen. The servants were not allowed to use the main stairs, but were restricted to the much narrower and steeper servants' stairs.
Tourism[edit]
Neuschwanstein welcomes almost 1.5 million visitors per year making it one of the most popular tourist destinations in Europe.[3][57] For security reasons the palace can only be visited during a 35-minute guided tour, and no photography is allowed inside the castle. There are also special guided tours that focus on specific topics. In the peak season from June until August, Neuschwanstein has as many as 6,000 visitors per day, and guests without advance reservation may have to wait several hours. Those without tickets may still walk the long driveway from the base to the top of the mountain and visit the grounds and courtyard without a ticket, but will not be admitted to the interior of the castle. Ticket sales are processed exclusively via the ticket centre in Hohenschwangau.[58] As of 2008, the total number of visitors was more than 60 million.[2] In 2004, the revenues were booked as €6.5 million.[1]
In culture, art, and science[edit]
Neuschwanstein is a global symbol of the era of Romanticism. The palace has appeared prominently in several movies such as Helmut Käutner's Ludwig II (1955) and Luchino Visconti's Ludwig (1972), both biopics about the King; the musical Chitty Chitty Bang Bang (1968) and the war drama The Great Escape (1963). It served as the inspiration for Disneyland's Sleeping Beauty Castle, Cameran Palace in Lucario and The Mystery of Mew, and later similar structures.[59][60] It is also visited by the character Grace Nakimura alongside Herrenchiemsee in the game The Beast Within: A Gabriel Knight Mystery (1996).
In 1977, Neuschwanstein Castle became the motif of a West German definitive stamp, and it appeared on a €2 commemorative coin for the German Bundesländer series in 2012. In 2007, it was a finalist in the widely publicised on-line selection of the New Seven Wonders of the World.[61]
A meteorite that reached Earth spectacularly on 6 April 2002, at the Austrian border near Hohenschwangau was named Neuschwanstein after the palace. Three fragments were found: Neuschwanstein I (1.75 kg (3.9 lb), found July 2002) and Neuschwanstein II (1.63 kg (3.6 lb), found May 2003) on the German side, and Neuschwanstein III (2.84 kg (6.3 lb), found June 2003) on the Austrian side near Reutte.[62] The meteorite is classified as an enstatite chondrite with unusually large proportions of pure iron (29%), enstatite and the extremely rare mineral sinoite (Si2N2O).[63]
World Heritage candidature[edit]
Since 2015, Neuschwanstein and Ludwig's Linderhof and Herrenchiemsee palaces are on the German tentative list for a future designation as UNESCO World Heritage Sites. A joint candidature with other representative palaces of the romantic historicism is discussed (including Schwerin Palace, for example).[64]
en.wikipedia.org/wiki/Neuschwanstein_Castle
Synopsis[edit]
Background[edit]
In Eisenach, Germany, in the early 13th century, the landgraves of the Thuringian Valley ruled the area of Germany around the Wartburg. They were great patrons of the arts, particularly music and poetry, holding contests between the minnesingers at the Wartburg. Across the valley towered the Venusberg, in whose interior, according to legend, dwelt Holda, the Goddess of Spring. In time, Holda became identified with Venus, the pagan Goddess of Love, whose grotto was the home of sirens and nymphs. It was said that the Goddess would lure the Wartburg minnesinger-knights to her lair where her beauty would captivate them. The minnesinger-knight Heinrich von Ofterdingen, known as Tannhäuser, left the court of the Landgrave of Thuringia a year ago after a disagreement with his fellow knights. Since then he has been held as a willing captive through his love for Venus, in her grotto in the Venusberg.[27][incomplete short citation][17]
Overture[edit]
The substantial overture commences with the theme of the 'Pilgrim's Chorus' from Act 3, Scene 1, and also includes elements of the 'Venusberg' music from Act 1, Scene 1. The overture is frequently performed as a separate item in orchestral concerts, the first such performance having been given by Felix Mendelssohn conducting the Leipzig Gewandhaus Orchestra in February 1846.[28] Wagner later gave the opinion that perhaps it would be better to cut the overture at opera performances to the Pilgrim's Chorus alone – "the remainder – in the fortunate event of its being understood – is, as a prelude to the drama, too much; in the opposite event, too little."[29] In the original, "Dresden" version, the overture comes to a traditional concert close (the version heard in concert performances). For the "Paris" version the music leads directly into the first scene, without pausing.
Act 1[edit]
The Venusberg, (the Hörselberg of "Frau Holda" in Thuringia, in the vicinity of Eisenach), and a valley between the Venusberg and Wartburg
Scene 1. Wagner's stage directions state: "The stage represents the interior of the Venusberg...In the distant background is a bluish lake; in it one sees the bathing figures of naiads; on its elevated banks are sirens. In the extreme left foreground lies Venus bearing the head of the half kneeling Tannhäuser in her lap. The whole cave is illuminated by rosy light. – A group of dancing nymphs appears, joined gradually by members of loving couples from the cave. – A train of Bacchantes comes from the background in wild dance... – The ever-wilder dance answers as in echo the Chorus of Sirens": "Naht euch dem Strande" (Come to the shore).[30] In the "Paris" version this orgiastic ballet is greatly extended.
Scene 2. Following the orgy of the ballet, Tannhäuser's desires are finally satiated, and he longs for freedom, spring and the sound of church bells. He takes up his harp and pays homage to the goddess in a passionate love song, "Dir töne Lob!" (Let your praises be heard), which he ends with an earnest plea to be allowed to depart, "Aus deinem Reiche, muss ich fliehn! O Königin! Göttin! Lass mich ziehn!" (From your kingdom must I flee! O Queen! O Goddess, set me free). Surprised, Venus offers him further charms, but eventually his repeated pleas arouse her fury and she curses his desire for salvation. (In the "Paris" version Venus's inveighing against Tannhäuser is significantly expanded).[31] Eventually Tannhäuser declares: "Mein Heil ruht in Maria" (My salvation rests in Mary). These words break the unholy spell. Venus and the Venusberg disappear.
Scene 3. According to Wagner's stage directions, "Tannhäuser...finds himself a beautiful valley… To the left one sees the Hörselberg. To the right...a mountain path from the direction of the Wartburg ...; in the foreground, led to by a low promontory, an image of the Virgin Mary – From above left one hears the ringing of herder’s bells; on a high projection sits a young shepherd with pipes facing the valley".[32] It is May. The shepherd sings an ode to the pagan goddess Holda, "Frau Holda kam aus dem Berg hervor" (Lady Holda, come forth from the hill). A hymn "Zu dir wall ich, mein Jesus Christ" (To thee I turn, my Jesus Christ) can be heard, as Pilgrims are seen approaching from the Wartburg, and the shepherd stops playing. The pilgrims pass Tannhäuser as he stands motionless, and then, praising God, ("Allmächt'ger, dir sei Preis!" (Almighty God, to you be praise!)) he sinks to his knees, overcome with gratitude. At that moment the sound of hunting-horns can be heard, drawing ever nearer.
Scene 4. The Landgrave's hunting party appears. The minnesingers (Wolfram, Walther, Biterolf, Reinmar, and Heinrich) recognise Tannhäuser, still deep in prayer, and greet him ("Heinrich! Heinrich! Seh ich recht?" (Heinrich! Heinrich! Do I see right?)) cautiously, recalling past feuds. They question him about his recent whereabouts, to which he gives vague answers. The minnesingers urge Tannhäuser to rejoin them, which he declines until Wolfram mentions Elisabeth, the Landgrave's niece, "Bleib bei Elisabeth!" (Stay, for Elisabeth!). Tannhäuser is visibly moved, "Elisabeth! O Macht des Himmels, rufst du den süssen Namen mir?" (Elisabeth! O might of heaven, do you cry out the sweet name to me?). The minnesingers explain to Tannhäuser how he had enchanted Elisabeth, but when he had left she withdrew from their company and lost interest in music, expressing the hope that his return will also bring her back, "Auf's Neue leuchte uns ihr Stern!" (Let her star once more shine upon us). Tannhäuser begs them to lead him to her, "Zu ihr! Zu ihr!" (To her! To her!). The rest of the hunting party gathers, blowing horns.
Act 2[edit]
The Wartburg in Eisenach
The minnesingers' hall in the Wartburg castle
Introduction – Scene 1. Elisabeth enters, joyfully. She sings, to the hall, of how she has been beset by sadness since Tannhäuser's departure but now lives in hope that his songs will revive both of them, "Dich, teure Halle, grüss ich wieder" (Dear hall, I greet thee once again). Wolfram leads Tannhäuser into the hall.
Scene 2. Tannhäuser flings himself at Elisabeth's feet. He exclaims "O Fürstin!" (O Princess!). At first, seemingly confused, she questions him about where he has been, which he avoids answering. She then greets him joyfully ("Ich preise dieses Wunder aus meines Herzens Tiefe!" (I praise this miracle from my heart's depths!)), and they join in a duet, "Gepriesen sei die Stunde" (Praise be to this hour). Tannhäuser then leaves with Wolfram.
Scene 3. The Landgrave enters, and he and Elisabeth embrace. The Landgrave sings of his joy, "Dich treff ich hier in dieser Halle" (Do I find you in this hall) at her recovery and announces the upcoming song contest, at which she will preside, "dass du des Festes Fürstin seist" (that you will be the Princess of the Festival).
Scene 4 and Sängerkrieg (Song Contest). Elisabeth and the Landgrave watch the guests arrive. The guests assemble greeting the Landgrave and singing "Freudig begrüssen wir edle Halle" (With joy we greet the noble hall), take their places in a semicircle, with Elisabeth and the Landgrave in the seats of honour in the foreground. The Landgrave announces the contest and the theme, which shall be "Könnt ihr der Liebe Wesen mir ergründen?" (Can you explain the nature of Love?), and that the prize will be whatever the winner asks of Elisabeth. The knights place their names in a cup from which Elisabeth draws the first singer, Wolfram. Wolfram sings a trite song of courtly love and is applauded, but Tannhäuser chides him for his lack of passion. There is consternation, and once again Elisabeth appears confused, torn between rapture and anxiety. Biterolf accuses him of blasphemy and speaks of "Frauenehr und hohe Tugend" (women's virtue and honour). The knights draw their swords as Tannhäuser mocks Biterolf, but the Landgrave intervenes to restore order. However, Tannhäuser, as if in a trance, rises to his feet and sings a song of ecstatic love to Venus, "Dir Göttin der Liebe, soll mein Lied ertönen" (To thee, Goddess of Love, should my song resound). There is general horror as it is realised he has been in the Venusberg; the women, apart from Elisabeth, flee. She appears pale and shocked, while the knights and the Landgrave gather together and condemn Tannhäuser to death. Only Elisabeth, shielding him with her body, saves him, "Haltet ein!" (Stop!). She states that God's will is that a sinner shall achieve salvation through atonement. Tannhäuser collapses as all hail Elisabeth as an angel, "Ein Engel stieg aus lichtem Äther" (An angel rose out of the bright ether). He promises to seek atonement, the Landgrave exiles him and orders him to join another younger band of pilgrims then assembling. All depart, crying Nach Rom! (To Rome!).
In the "Paris" version, the song contest is somewhat shortened, possibly because of the lack of suitable soloists for the Paris production.[citation needed]
Act 3[edit]
The valley of the Wartburg, in autumn. Elisabeth is kneeling, praying before the Virgin as Wolfram comes down the path and notices her
Scene 1. Orchestral music describes the pilgrimage of Tannhäuser. It is evening. Wolfram muses on Elisabeth's sorrow during Tannhäuser's second absence, "Wohl wusst' ich hier sie im Gebet zu finden" (I knew well I might find her here in prayer) and her longing for the return of the pilgrims, and expresses concerns that he may not have been absolved. As he does so he hears a pilgrims' prayer in the distance, "Beglückt darf nun dich, O Heimat, ich schauen" (Joyfully may I now you, O homeland, behold). Elisabeth rises and she and Wolfram listen to the hymn, watching the pilgrims approach and pass by. She anxiously searches the procession, but in vain, realising sorrowfully he is not amongst them, "Er kehret nicht züruck!" (He has not returned). She again kneels with a prayer to the Virgin that appears to foretell her death, "Allmächt'ge Jungfrau! Hör mein Flehen" (Almighty Virgin, hear my plea!). On rising she sees Wolfram but motions him not to speak. He offers to escort her back to the Wartburg, but she again motions him to be still, and gestures that she is grateful for his devotion but her path leads to heaven. She slowly makes her way up the path alone.
Scene 2. Wolfram, left alone as darkness draws on and the stars appear, begins to play and sings a hymn to the evening star that also hints at Elisabeth's approaching death, "Wie Todesahnung Dämmrung deckt die Lande...O du mein holder Abendstern" (Like a premonition of death the twilight shrouds the earth... O thou my fair evening star).
Scene 3. It is now night. Tannhäuser appears, ragged, pale and haggard, walking feebly leaning on his staff. Wolfram suddenly recognises Tannhäuser, and startled challenges him, since he is exiled. To Wolfram's horror, Tannhäuser explains he is once again seeking the company of Venus. Wolfram tries to restrain him, at the same time expressing compassion and begging him to tell the story of his pilgrimage. Tannhäuser urges Wolfram to listen to his story, "Nun denn, hör an! Du, Wolfram, du sollst es erfahren" (Now then, listen! You, Wolfram, shall learn all that has passed). Tannhäuser sings of his penitence and suffering, all the time thinking of Elisabeth's gesture and pain, "Inbrunst im Herzen, wie kein Büsser noch" (With a flame in my heart, such as no penitent has known). He explains how he reached Rome, and the "Heiligtumes Schwelle" (Holy shrine), and witnessed thousands of pilgrims being absolved. Finally he approaches "ihn, durch den sich Gott verkündigt'" (he, through whom God speaks)[a] and tells his story. However, rather than finding absolution, he is cursed, "bist nun ewig du verdammt!" (you are forever damned!), and is told by the pope that "Wie dieser Stab in meiner Hand, nie mehr sich schmückt mit frischem Grün, kann aus der Hölle heissem Brand, Erlösung nimmer dir erblühn!" (As this staff in my hand, no more shall bear fresh leaves, from the hot fires of hell, salvation never shall bloom for thee). Whereupon, absolutely crushed, he fled, seeking his former source of bliss.
Having completed his tale, Tannhäuser calls out to Venus to take him back, "Zu dir, Frau Venus, kehr ich wieder" (To you, Lady Venus, I return). The two men struggle as a faint image of dancing becomes apparent. As Tannhäuser repeatedly calls on Venus, she suddenly appears and welcomes him back, "Willkommen, ungetreuer Mann!" (Welcome, faithless man!). As Venus continues to beckon, "Zu mir! Zu mir!" (To me!, To me!), in desperation, Wolfram suddenly remembers there is one word that can change Tannhäuser's heart, and exclaims "Elisabeth!" Tannhäuser, as if frozen in time, repeats the name. As he does so, torches are seen, and a funeral hymn is heard approaching, "Der Seele Heil, die nun entflohn" (Hail, the soul that now is flown). Wolfram realises it must be Elisabeth's body that is being borne, and that in her death lies Tannhäuser's redemption, "Heinrich, du bist erlöst!" (Heinrich, you are saved). Venus cries out, "Weh! Mir verloren" (Alas! Lost to me!) and vanishes with her kingdom. As dawn breaks the procession appears bearing Elisabeth's body on a bier. Wolfram beckons to them to set it down, and as Tannhäuser bends over the body uttering, "Heilige Elisabeth, bitte für mich!" (Holy Elisabeth!, pray for me!) he dies. As the growing light bathes the scene the younger pilgrims arrive bearing the pope's staff sprouting new leaves, and proclaiming a miracle, "Heil! Heil! Der Gnade Wunder Heil!" (Hail!, Hail! To this miracle of grace, Hail!). All then sing "Der Gnade Heil ist dem Büsser beschieden, er geht nun ein in der Seligen Frieden!" (The Holy Grace of God is to the penitent given, who now enters into the joy of Heaven!).[27][incomplete short citation][25][30]
After Wagner[edit]
Productions[edit]
Wagner died in 1883. The first production of the opera at Wagner's Bayreuth Festspielhaus (originally constructed for the performance of his Ring Cycle), was undertaken under the supervision of Cosima in 1891, and adhered closely to the 'Vienna' version. Later performances at Bayreuth included one conducted by Richard Strauss (1894), and one where the Bacchanal was choreographed by Isadora Duncan (1904).[33] Duncan envisaged the Bacchanal as a fantasy of Tannhäuser's fevered brain, as Wagner had written to Mathilde Wesendonck in 1860.[34] Arturo Toscanini conducted the opera at Bayreuth in the 1930/31 season.[35][incomplete short citation]
In the words of the Wagner scholar Thomas S. Grey, "The Bacchanal remained a defining focus of many ...productions, as a proving ground for changing conceptions of the psychosexual symbolism of the Venusberg." Productions including those of Götz Friedrich at Bayreuth (1972) and Otto Schenk at the Metropolitan Opera, New York, (1977) "routinely offer quantities of simulated copulation and post-coital langour, for which the Paris score offers ample encouragement".[33] A Munich production (1994) included as part of Tannhäuser's fantasies "creatures out of Hieronymus Bosch crawl[ing] around the oblivious protagonist".[36]
The Operabase website indicates that in the two calendar years 2014/2015, there were 163 performances of 41 productions of Tannhäuser in 30 cities throughout the world.[37]
Literature[edit]
Many scholars and writers on opera have advanced theories to explain the motives and behaviour of the characters,[9] including Jungian psychoanalysis,[1] in particular as regards Tannhäuser's apparently self-destructive behaviour. In 2014 an analysis suggested that his apparently inconsistent behaviour, when analysed by game theory, is actually consistent with a redemption strategy. Only by public disclosure can Tannhäuser force a resolution of his inner conflict.[38]
夢幻蝶影 蝶戀花
Dream of BUTTERFLY WITH
The Great Mormon (Papilio memnon) is a large butterfly that belongs to the swallowtail family and is found in southern Asia. It is widely distributed and has thirteen subspecies. The female is polymorphic and with mimetic forms.
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Papilio memnon, the great Mormon, is a large butterfly native to southern Asia that belongs to the swallowtail family. It is widely distributed and has thirteen subspecies. The female is polymorphic and with mimetic forms. Its range includes north-eastern India (including Sikkim, Assam and Nagaland), Nepal, Bangladesh, Myanmar, Nicobar Islands, Andaman Islands (stragglers only), western, southern and eastern China (including Hainan), Taiwan southern Japan, Ryukyu Islands, Thailand, Laos, Vietnam, Kampuchea, Malaysia and Indonesia (Sumatra, Mentawai Islands, Nias, Batu, Simeulue, Bangka, Java, Kalimantan and the Lesser Sunda Islands).The butterfly is large with a 120 to 150 millimetres (4.7 to 5.9 in) span. It has four male and many female forms, the females being highly polymorphic and many of them being mimics of unpalatable butterflies. This species has been studied extensively for understanding the genetic basis for polymorphy and Batesian mimicry. As many as twenty-six female forms are reported
Photographed at Los Altos, California - Sitting, no cover
=> Please click on the image to see the largest size. <=
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From Wikipedia: The lesser goldfinch (Spinus psaltria) is a very small songbird of the Americas. Together with its relatives the American goldfinch and Lawrence's goldfinch, it forms the American goldfinches clade in the genus Spinus sensu stricto.
The American goldfinches can be distinguished by the males having a black (rarely green) forehead, whereas the latter is (like the rest of the face) red or yellow in the European goldfinch and its relatives. North American males are markedly polymorphic and five subspecies are often named; at least two of them seem to represent a less-progressed stage in evolution, however.
This petite species is not only the smallest North American Spinus finch, it may be the smallest true finch in the world. Some sources list more subtropical Spinus species as slightly smaller on average, including the Andean siskin.[4] This species ranges from 9 to 12 cm (3.5 to 4.7 in) in length and can weigh from 8 to 11.5 g (0.28 to 0.41 oz). Among standard measurements, the wing chord is 5.5 to 7 cm (2.2 to 2.8 in), the tail is 3.9 to 4.7 cm (1.5 to 1.9 in), the bill is 0.9 to 1.1 cm (0.35 to 0.43 in) and the tarsus is 1.1 to 1.2 cm (0.43 to 0.47 in). There is a slight NW-SE cline in size, with the largest birds from Mexico and further south being up to one-fifth larger than the smallest from the extreme northwest of its range; this effect is more pronounced in females. There is also considerable variation in the amount of black on head and back in males, and thus three subspecies have been proposed. But this variation, too, seems to be simple and clinal changes in allele frequency, and thus the "subspecies" might be better considered morphs or geographic forms.
Males are easily recognized by their bright yellow underparts and big white patches in the tail (outer rectrices) and on the wings (the base of the primaries). They range from having solid black from the back to the upper head including the ear-coverts to having these regions medium green; each of the back, crown and ear regions varies in darkness rather independently though, as a rule, the ears are not darker than the rest. In most of the range, dark psaltria birds (Arkansas goldfinch) predominate. The light birds are termed hesperophilus (green-backed goldfinch) and are most common in the far western U.S. and northwestern Mexico.
Females' and immatures' upperparts are more or less grayish olive-green; their underparts are yellowish, buffier in immatures. They have only a narrow strip of white on the wings (with other white markings in some forms) and little or no white on the tail. They are best distinguished from other members of the genus by the combination of small size, upperparts without white or yellow, and dark gray bill. In all plumages, this bird can easily be taken for a New World warbler if the typical finch bill is not seen well.
Like other goldfinches, it has an undulating flight in which it frequently gives a call: in this case, a harsh chig chig chig. Another distinctive call is a very high-pitched, drawn-out whistle, often rising from one level pitch to another (teeeyeee) or falling (teeeyooo). The song is a prolonged warble or twitter, more phrased than that of the American goldfinch, often incorporating imitations of other species.
Distribution and ecology:
Lesser Goldfinch Landing, Santa Fe.jpg
This American goldfinch ranges from the southwestern United States (near the coast, as far north as extreme southwestern Washington) to Venezuela and Peru. It migrates from the colder parts of its U.S. range.
The lesser goldfinch often occurs in flocks or at least loose associations. It utilizes almost any habitat with trees or shrubs except for dense forest, and is common and conspicuous in many areas, often coming near houses. It is common at feeders in the Southwest United States and will come almost anywhere with thistle sock feeders. Flocks of at least six birds will often be seen at feeders. It feeds mostly on tree buds and weed seeds; geophagy has been observed in this species.
The nesting season is in summer in the temperate parts of its range; in the tropics it apparently breeds all-year round, perhaps less often in September/October. It lays three or four bluish white eggs in a cup nest made of fine plant materials such as lichens, rootlets, and strips of bark, placed in a bush or at low or middle levels in a tree.
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POLYMORPHIC SEQUENCE OF THE REALITY GRID / THE FINAL / CHRISTELLE GEISER & AEON VON ZARK / NAKED EYE PROJECT BIENNE / ALTERED STATE SERIE / THE WEIRD DREAM / PORTRAIT.
The Great Mormon butterfly, Papilio memnon, is a large swallowtail butterfly (family Papilionidae) with wingspan up to 135mm, native to and common throughout Southeast Asia. Larvae feed on citrus species, particularly pomelo (citrus grandis) and common lime (citrus aurantifolia) which are broadly cultivated across Asia. Adults lay single eggs on the underside of leaves, which hatch in three days. For the first four instars, caterpillars resemble bird droppings. Pupation occurs after about 2.5 weeks. Adults inhabit forest clearings and disturbed areas, eating nectar from a wide variety of flowering species.
Alfred Russel Wallace first described the remarkable polymorphic nature of P. memnon, which exists as 13 subspecies and in addition, females show a large diversity of morphological and coloration forms, many of which are mimetic of other unpalatable papilionid species (Batesian mimicry). Female morphological variations include: presence or absence of tails, hindwing pattern, forewing pattern, color of the basal triangle on the forewing, and abdomen color. Extensive study of this species has contributed insight into the genetic determination of mimicry. These studies give classic evidence for existence of a “supergene” complex, which slowly built up over the course of evolution allowing butterfly species to mimic their models very accurately. This complex includes multiple linked genes that control the morphological variations listed above via genetic crossover, rather than these traits diversifying individually by point mutations, or other genetic mechanism.
কালিম । Common Mormon (Papilio polytes) - Male
A common species of swallowtail butterfly widely distributed across Asia. Seen round the year throughout India from plains up to 2000m. This butterfly is known for the mimicry displayed by the numerous polymorphic forms of its females. These are as follows: cyrus, stichius, romulus.
Family: Papilionidae
Butterfly Garden, Garpanchakot Forest, Purulia District
Butterflies of Bengal, India
Cavansite, whose name is derived from its chemical composition, calcium vanadium silicate, is a deep blue hydrous calcium vanadium phyllosilicate mineral, occurring as a secondary mineral in basaltic and andesitic rocks along with a variety of zeolite minerals. Discovered in 1967 in Malheur County, Oregon, cavansite is a relatively rare mineral. It is polymorphic with the even rarer mineral, pentagonite. It is most frequently found in Pune, India and in the Deccan Traps, a large igneous province.
Doris longwing
The Laparus doris is a polymorphic butterflies that comes in several different forms, for the most part in three colors: red, blue and green.
2011 Butterflies of Brazil Krohn Conservatory Cincinnati OH
*** check out the entire set "2011 Butterflies of Brazil"
Photographed while walking at San Antonio Open Space Preserve, Los Altos, California
Please click on the photo or press the L key to view the larger size
This beautiful Red-tailed Hawk was perched on a horizontal branch, no more than 50 feet from a heavily-used trail that winds up the hillside from the parking area. Many hikers and runners passed this hawk in both directions without noticing the nearby hawk. The hawk itself was constantly moving its head about as it was searching for prey and in this photo was looking in my general direction at movement off to my right.
Canon 7D Mark II. f/6.3 1/640 ISO 400
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From Wikipedia: The red-tailed hawk (Buteo jamaicensis) is a bird of prey that breeds throughout most of North America, from the interior of Alaska and northern Canada to as far south as Panama and the West Indies. It is one of the most common members within the genus of Buteo in North America or worldwide. The red-tailed hawk is one of three species colloquially known in the United States as the "chickenhawk," though it rarely preys on standard-sized chickens. The bird is sometimes also referred to as the red-tail for short, when the meaning is clear in context.
Red-tailed hawks can acclimate to all the biomes within their range, occurring on the edges of non-ideal habitats such as dense forests and sandy deserts. The red-tailed hawk occupies a wide range of habitats and altitudes including deserts, grasslands, coniferous and deciduous forests, agricultural fields and urban areas. Its latitudinal limits fall around the tree line in the Arctic and the species is absent from the high Arctic. It is legally protected in Canada, Mexico and the United States by the Migratory Bird Treaty Act.
The 14 recognized subspecies vary in appearance and range, varying most often in color, and in the west of North America, red-tails are particularly often strongly polymorphic, with individuals ranging from almost white to nearly all black. The subspecies Harlan's hawk (B. j. harlani) is sometimes considered a separate species (B. harlani). The red-tailed hawk is one of the largest members of the genus Buteo, typically weighing from 690 to 1,600 g (1.5 to 3.5 lb) and measuring 45–65 cm (18–26 in) in length, with a wingspan from 110–141 cm (3 ft 7 in–4 ft 8 in). This species displays sexual dimorphism in size, with females averaging about 25% heavier than males.
The diet of red-tailed hawks is highly variable and reflects their status as opportunistic generalist, but in North America, it is most often a predator of small mammals such as rodents. Prey that is terrestrial and diurnal is preferred so types such as ground squirrels are preferential where they naturally occur. Large numbers of birds and reptiles can occur in the diet in several areas and can even be the primary foods. Meanwhile, amphibians, fish and invertebrates can seem rare in the hawk’s regular diet; however, they are not infrequently taken by immature hawks.
Red-tailed hawks may survive on islands absent of native mammals on diets variously including invertebrates such as crabs, or lizards and birds. Like many Buteo, they hunt from a perch most often but can vary their hunting techniques where prey and habitat demand it. Because they are so common and easily trained as capable hunters, the majority of hawks captured for falconry in the United States are red-tails. Falconers are permitted to take only passage hawks (which have left the nest, are on their own, but are less than a year old) so as to not affect the breeding population. Adults, which may be breeding or rearing chicks, may not be taken for falconry purposes and it is illegal to do so. Passage red-tailed hawks are also preferred by falconers because these younger birds have not yet developed the adult behaviors which would make them more difficult to train.
Description:
Red-tailed hawk plumage can be variable, depending on the subspecies and the region. These color variations are morphs, and are not related to molting. The western North American population, B. j. calurus, is the most variable subspecies and has three main color morphs: light, dark, and intermediate or rufous. The dark and intermediate morphs constitute 10–20% of the population in the western United States but seem to constitute only 1-2% of B. j. calurus in western Canada. A whitish underbelly with a dark brown band across the belly, formed by horizontal streaks in feather patterning, is present in most color variations. This feature is variable in eastern hawks and generally absent in some light subspecies (i.e. B. j. fuertesi).
Most adult red-tails have a dark brown nape and upper head which gives them a somewhat hooded appearance, while the throat can variably present a lighter brown “necklace”. Especially in younger birds, the underside may be otherwise covered with dark brown spotting and some adults may too manifest this stippling. The back is usually a slightly darker brown than elsewhere with paler scapular feathers, ranging from tawny to white, forming a variable imperfect “V” on the back. The tail of most adults, which of course gives this species its name, is rufous brick-red above with a variably sized black subterminal band and generally appears light buff-orange from below. In comparison, the typical pale immatures (i.e. less than two years old) typically have a mildly paler headed and tend to show a darker back than adults with more apparent pale wing feather edges above (for descriptions of dark morph juveniles from B. j. calurus, which is also generally apt for description of rare dark morphs of other races, see under that subspecies description). In immature red-tailed hawks of all hues, the tail is a light brown above with numerous small dark brown bars of roughly equal width, but these tend to be much broader on dark morph birds.
Even in young red-tails, the tail may be a somewhat rufous tinge of brown. The bill is relatively short and dark, in the hooked shape characteristic of raptors, and the head can sometimes appear small in size against the thick body frame. The cere, the legs, and the feet of the red-tailed hawk are all yellow, as is the hue of bare parts in many accipitrids of different lineages. Immature birds can be readily identified at close range by their yellowish irises. As the bird attains full maturity over the course of 3–4 years, the iris slowly darkens into a reddish-brown hue, which is the adult eye-color in all races. Seen in flight, adults usually have dark brown along the lower edge of the wings, against a mostly pale wing, which bares light brownish barring. Individually, the underwing coverts can range from all dark to off-whitish (most often more heavily streaked with brown) which contrasts with a distinctive black patagium marking. The wing coloring of adults and immatures is similar but for typical pale morph immatures having somewhat heavier brownish markings.
Though the markings and hue vary across the subspecies, the basic appearance of the red-tailed hawk is relatively consistent. Overall, this species is blocky and broad in shape, often appearing (and being) heavier than other Buteos of similar length. They are the heaviest Buteos on average in eastern North America, albeit scarcely ahead of the larger winged rough-legged buzzard (Buteo lagopus), and second only in size in the west to the ferruginous hawk (Buteo regalis). Red-tailed hawks may be anywhere from the seventh to the ninth heaviest Buteo in the world depending on what figures are used. However, in the northwestern United States, ferruginous hawk females are 35% heavier than female red-tails from the same area. On average, western red-tailed hawks are relatively longer winged and lankier proportioned but are slightly less stocky, compact and heavy than eastern red-tailed hawks in North America. Eastern hawks may also have mildly larger talons and bills than western ones. Based on comparisons of morphology and function amongst all accipitrids, these features imply that western red-tails may need to vary their hunting more frequently to on the wing as the habitat diversifies to more open situations and presumably would hunt more variable and faster prey, whereas the birds of the east, which was historically well-wooded, are more dedicated perch hunters and can take somewhat larger prey but are likely more dedicated mammal hunters. In terms of size variation, red-tailed hawks run almost contrary to Bergmann's rule (i.e. that northern animals should be larger in relation than those closer to the Equator within a species) as one of the northernmost subspecies, B. j. alascensis, is the second smallest race based on linear dimensions and that two of the most southerly occurring races in the United States, B. j. fuertesi and B. j. umbrinus, respectively, are the largest proportioned of all red-tailed hawks. Red-tailed hawks tend have a relatively short but broad tails and thick, chunky wings. Although often described as long winged, the proportional size of the wings is quite small and red-tails have high wing loading for a buteonine hawk. For comparison, two other widespread Buteo hawks in North America were found to weigh: 30 g (1.1 oz) for every square centimeter of wing area in the rough-legged buzzard (Buteo lagopus) and 44 g (1.6 oz) per square cm in the red-shouldered hawk (Buteo lineatus). In contrast, the red-tailed hawk weighed considerably more for their wing area: 199 g (7.0 oz) per square cm.
As is the case with many raptors, the red-tailed hawk displays sexual dimorphism in size, as females are up to 25% larger than males. As is typical in large raptors, frequently reported mean body mass for Red-tailed Hawks are somewhat higher than expansive research reveals. Part of this weight variation is seasonal fluctuations, hawks tending to be heavier in winter than during migration or especially during the trying summer breeding season, and also due to clinal variation. Furthermore, immature hawks are usually lighter in mass than their adult counterparts despite averaging somewhat longer winged and tailed. Male red-tailed hawks may weigh from 690 to 1,300 g (1.52 to 2.87 lb) and females may weigh between 801 and 1,723 g (1.766 and 3.799 lb) (the lowest figure from a migrating female immature from Goshute Mountains, Nevada, the highest from a wintering female in Wisconsin). Some sources claim the largest females can weigh up to 2,000 g (4.4 lb) but whether this is in reference to wild hawks (as opposed to those in captivity or used for falconry) is not clear.[24] The largest known survey of body mass in red-tailed hawks is still credited to Craighead & Craighead (1956), who found 100 males to average 1,028 g (2.266 lb) and 108 females to average 1,244 g (2.743 lb). However, these figures were apparently taken from labels on museum specimens, apparently from natural history collections in Wisconsin and Pennsylvania, without note to the region, age or subspecies of the specimens. However, 16 sources ranging in sample size from the aforementioned 208 specimens to only four hawks in Puerto Rico (with 9 of the 16 studies of migrating red-tails), showed that males weigh a mean of 860.2 g (1.896 lb) and females weigh a mean of 1,036.2 g (2.284 lb), about 15% lighter than prior species-wide published weights. Within the continental United States, average weights of males can range from 840.8 g (1.854 lb) (for migrating males in Chelan County, Washington) to 1,031 g (2.273 lb) (for male hawks found dead in Massachusetts) and females ranged from 1,057.9 g (2.332 lb) (migrants in the Goshutes) to 1,373 g (3.027 lb) (for females diagnosed as B. j. borealis in western Kansas). Size variation in body mass reveals that the red-tailed hawks typically varies only a modest amount and that size differences are geographically inconsistent. Racial variation in average weights of great horned owls (Bubo virginianus) show that mean body mass is nearly twice (the heaviest race is about 36% heavier than the lightest known race on average) as variable as that of the hawk (where the heaviest race is only just over 18% heavier on average than the lightest). Also, great horned owls correspond well at the species level with Bergmann’s rule.
Male red-tailed hawks can reportedly measure 45 to 60 cm (18 to 24 in) in total length, females measuring 48 to 65 cm (19 to 26 in) long. The wingspan typically can range from 105 to 141 cm (3 ft 5 in to 4 ft 8 in), although the largest females may possible span up to 147 cm (4 ft 10 in). In the standard scientific method of measuring wing size, the wing chord is 325.1–444.5 mm (12.80–17.50 in) long. The tail measures 188 to 258.7 mm (7.40 to 10.19 in) in length. The exposed culmen was reported to range from 21.7 to 30.2 mm (0.85 to 1.19 in) and the tarsus averaged 74.7–95.8 mm (2.94–3.77 in) across the races. The middle toe (excluding talon) can range from 38.3 to 53.8 mm (1.51 to 2.12 in), with the hallux-claw (the talon of the rear toe, which has evolved to be the largest in accipitrids) measuring from 24.1 to 33.6 mm (0.95 to 1.32 in) in length.
Identification:
Although they overlap in range with most other American diurnal raptors, identifying most mature red-tailed hawks to species is relatively straightforward, particularly if viewing a typical adult at a reasonable distance. The red-tailed hawk is the only North American hawk with a rufous tail and a blackish patagium marking on the leading edge of its wing (which is obscured only on dark morph adults and Harlan’s hawks by similarly dark colored feathers).
Other larger adult Buteo in North America usually have obvious distinct markings that are absent in red-tails, whether the rufous-brown “beard” of Swainson's hawks (Buteo swainsonii) or the colorful rufous belly and shoulder markings and striking black-and-white mantle of red-shouldered hawks (also the small “windows” seen at the end of their primaries). In perched individuals, even as silhouettes, the shape of large Buteos may be distinctive, such as the wingtips overhanging the tail in several other species, but not in red-tails. North American Buteos range from the dainty, compact builds of much smaller Buteos, such as broad-winged hawk (Buteo platypterus) to the heavyset, neckless look of ferruginous hawks or the rough-legged buzzard which has a compact, smaller appearance than a red-tail in perched birds due to its small bill, short neck and much shorter tarsus, while the opposite effect occurs in flying rough-legs with their much bigger wing area.
In flight, most other large North American Buteo are distinctly longer and slenderer winged than red-tailed hawks, with the much paler ferruginous hawk having peculiarly slender wings in relation to its massive, chunky body. Swainson's hawks are distinctly darker on the wing and ferruginous hawks are much paler winged than typical red-tailed hawks. Pale morph adult ferruginous hawk can show mildly tawny-pink (but never truly rufous) upper tail, and like red-tails tend to have dark markings on underwing-coverts and can have a dark belly band but compared to red-tailed hawks have a distinctly broader head, their remiges are much whiter looking with very small dark primary tips, they lack the red-tail’s diagnostic patagial marks and usually (but not always) also lack the dark subterminal tail-band, and ferruginous have a totally feathered tarsus. With its whitish head, the ferruginous hawk is most similar to Krider's red-tailed hawks, especially in immature plumage, but the larger hawk has broader head and narrower wing shape and the ferruginous immatures are paler underneath and on their legs. Several species share a belly band with the typical red-tailed hawk but they vary from subtle (as in the ferruginous hawk) to solid blackish, the latter in most light-morph rough-legged buzzards. More difficult to identify among adult red-tails are its darkest variations, as most species of Buteo in North America also have dark morphs. Western dark morph red-tails (i.e. calurus) adults, however, retain the typical distinctive brick-red tail which other species lack, which may stand out even more against the otherwise all chocolate brown-black bird. Standard pale juveniles when perched show a whitish patch in the outer half of the upper surface of the wing which other juvenile Buteo lack. The most difficult to identify stages and plumage types are dark morph juveniles, Harlan’s hawk and some Krider’s hawks (the latter mainly with typical ferruginous hawks as aforementioned). Some darker juveniles are similar enough to other Buteo juveniles that it has been stated that they "cannot be identified to species with any confidence under various field conditions." However, field identification techniques have advanced in the last few decades and most experienced hawk-watchers can distinguish even the most vexingly plumaged immature hawks, especially as the wing shapes of each species becomes apparent after seeing many. Harlan’s hawks are most similar to dark morph rough-legged buzzards and dark morph ferruginous hawks. Wing shape is the most reliable identification tool for distinguishing the Harlan’s from these, but also the pale streaking on the breast of Harlan’s, which tends to be conspicuous in most individuals, and is lacking in the other hawks. Also dark morph ferruginous hawks do not have the dark subterminal band of a Harlan’s hawk but do bear a black undertail covert lacking in Harlan’s.
AB2A8516-1_fCA2Flkrs
Littorina fabalis male. A large obvious penis, positioned at the posterior of the head on the right side, is present on mature males in all seasons, and can be exposed on dead or living specimens of both species.
Shell height 12.5mm. Llŷn, North Wales. September 2015.
FULL ACCOUNT BELOW
Sets of OTHER SPECIES:
www.flickr.com/photos/56388191@N08/collections/
PDF available at www.researchgate.net/publication/351037569_Differentiatin... .
Differentiating Littorina obtusata sensu stricto (Linnaeus, 1758)
from Littorina fabalis (Turton, 1825).
Ian F. Smith
Abstract
Adult males of Littorina obtusata sensu stricto and L. fabalis (synonym L. mariae) can be reliably differentiated by the forms of the penes on both extracted dead specimens and unsedated intact animals. Differentiation can often be made within a local population on the basis of shell form, size and colour after these features have been correlated with penis forms in a sample of specimens. The shell features can then be used to identify further local specimens, but cannot be relied on at other localities without confirming penis forms there.
Materials, methods, protocols for acquiring experience, environmental associations of phenotypes, and some sample sites are described and illustrated.
Contents
1. Preliminary check.
2. Taxonomic history.
3. Identification resources.
4. Penis examination methods.
5. Differences in penis morphology.
6. Differences in female anatomy.
7. Differences in shell morphology and colour.
7a. Shell colours.
7b. Shell sizes.
7c. Shell spires.
7d. Shell apertures.
8. Phenotypes of different wave exposures.
8a. Sheltered shores.
8b. Semi-exposed shores.
8c. More exposed shores.
9. Collecting and reliable recording.
10. Acknowledgements.
11. References and web links.
12. Glossary.
1. Preliminary check
Lacuna pallidula, detailed account below image flic.kr/p/hTnxJa , has a similar low spire, very similar spawn, and occurs with L. fabalis on Fucus serratus . It has a much thinner shell with widely open umbilicus, contrasting with the thick columella of L. fabalis.
2. Taxonomic history of L. obtusata and L. fabalis.
1758 to 1914: multiple species and varieties named.
1915 to 1965: all species, except Arctic varieties with spires, combined into one species referred to as Littorina obtusata (Linnaeus, 1758) or, mainly by British workers, as L. littoralis auct. For detail of historic nomenclature confusion see caption below image 1Dof flic.kr/p/QFCUWm .
1966 to 1988: gradual acceptance of two species, based on penis differences, named L. obtusata sensu stricto (Linnaeus, 1758) and L. fabalis (Turton, 1825), synonym L. mariae Sacchi & Rastelli, 1966. Arctic varieties with pointed spires were included in one or other of the pair on the basis of their penes.
In the following account, any use of “L. obtusata” in the now unaccepted sense of including both species has “sensu lato” or “s.l.” after it. The name with no addition, or sometimes for emphasis with “sensu stricto” or “s.s.”, refers to the now accepted segregated sense.
3. Identification resources
Differentiation of the two species by penis form was first published in 1966 by Sacchi & Rastelli, but did not appear in many identification guides until the late 1980s. Consequently, most pre 1988 works, apart from post 1966 specialist papers, are of limited use. Some later publications, such as Graham (1988), recognise the two species but provide insufficient information and images for discrimination of the many phenotypes. Hayward & Ryland (1990) and Hayward & Ryland (1995 & reprints to 2009), have images www.seawater.no/fauna/mollusca/fabalis.html that confuse www.ispotnature.org/node/646985 as the only L. mariae shell illustrated has features frequent on juveniles of both species, and the L. obtusata image has thick aperture walls more typical of L. fabalis.
Williams (1990) has useful information on forms and ecology. The authoritative volume Systematics and evolution of Littorina by Reid (1996) has fifty A4 pages of detailed description and comparison of all phenotypes and geographical variations of these two species. It is essential reading for those undertaking scientific study of Littorina species www.raysociety.org.uk/publications/zoology/the-systematic... .
4. Penis examination
“...only the characters of the adult reproductive system are unequivocal... . ...identifications using [shell features] should be confirmed by dissection before routine application in the field” (Reid, 1996). A rare ambiguity in penes linked to hybridization has been reported at one site in Portugal (Carvalho et al., 2016). An obvious penis, positioned at the posterior of the head on the right side 2Dof flic.kr/p/RcvnFf , is present on mature males in all seasons, and can be exposed on dead or living specimens.
Dead material
Specimens can be killed instantly by plunging into boiling water, or by leaving in a freezer overnight, or by gradually adding Magnesium chloride or Magnesium sulphate crystals to the water in a small container to first anaesthetize and then kill them. The only dissection required is to carefully crush the shell in a tabletop vice with any rubber protectors removed 3Dof flic.kr/p/QFCUmy , avoiding damage to the soft parts, and to pick off the broken fragmets. This method allows unrestricted view of all the necessary details and is quick and reliable, but requires killing of the specimen and destruction of the shell. Alternatively, if boiled for several minutes, the animal can be extracted with a pin from the unbroken shell. Though the penis as a whole is likely to contract on death, especially if boiled, and will be fixed in one of several possible expansions and positions 4Dof flic.kr/p/QFCUab , its glands are often distended and more easily discerned, and it is likely to resemble published images of dead specimens 4Dof flic.kr/p/QFCUab .
Living unsedated specimens
Penes can be examined and photographed on live specimens without harm or anaesthesia using the techniques described in Smith (2012 & 2016 flic.kr/s/aHskNP6GoL ). While live, the penis can be held in a variety of positions 5aDof flic.kr/p/Q2jb82 & 5bDof flic.kr/p/Rcvmo5 and will often be generally more expanded than on dead material, but the glands are often less prominent than on dissected penes and frequently hidden when held against the body. Time and patience will be required as this pair of species may be reluctant to extend from their shells, and several views or images may be needed to accumulate the necessary detail. Best results are obtained if the examination can be within 48 hours of capture. When not being examined, keep in a refrigerator at about 6°C in tightly closed plastic boxes dampened by, but not awash with, seawater. L. fabalis will extend when immersed and restrained 7Dof flic.kr/p/Q2jabc as described in Smith (2012 & 2016), but L. obtusata s.s. exposes itself more readily when damp than when submerged 8Dof flic.kr/p/RcvkCY & 9Dof flic.kr/p/Q2j9un .
5. Differences in penis morphology,
Mamilliform penial glands
L. fabalis has 3 to 17 large glands in a single row along 40% to 70% of the ventral edge of its penis 10Dof flic.kr/p/Rcvk2Y . The glands are usually visible on live unsedated specimens 7Dof flic.kr/p/Q2jabc .
L. obtusata has 10 to 54 closely packed small glands, in one or more irregular rows along 75% to 90% of the length of its penis 10Dof flic.kr/p/Rcvk2Y near its ventral edge and on the mesial face usually held against the body 9Dof flic.kr/p/Q2j9un ; sometimes multiple rows occupy 60% of the width of the penis. The small glands often do not protrude into view on live unsedated specimens unless the penis is twisted to expose the proximal face 11Dof flic.kr/p/RfXgF8 .
Filament (glandless tip of penis).
L. fabalis has long and vermiform filament. On dead or anaesthetized specimens filament is 30% to over 60% of total length of penis 10Dof flic.kr/p/Rcvk2Y . Growth is allometric; small individuals frequently have filaments 30% to 40%, while the largest usually have filaments of 50% to >60% of penis length 6Dof flic.kr/p/RTWK1i . Penis is motile and varying in length on any individual unsedated specimen 5bDof flic.kr/p/Rcvmo5 , and the appearance is affected by positioning/angle of view, so it is advisable to take several observations or photographs when examining live specimens.
L. obtusata has short triangular filament. On dead or anaesthetized specimens 10% to 25% of total length of penis 10Dof flic.kr/p/Rcvk2Y . On live unsedated specimens it may be difficult to discern the precise limit (distal gland) of the filament because the small glands are hidden. However, being short, wide and triangular, it is obviously different 12Dof flic.kr/p/RfXgop & 13Dof flic.kr/p/RfXg8z from the vermiform filament of L. fabalis.
Size of penis depends on the size of the individual male and is not diagnostic of species. On the sheltered Menai Strait, most adult male L. obtusata are larger, and so have larger penes, than most male L. fabalis. On more exposed shores in North Wales, the sizes of the two species can be almost equal.
6. Difference in female anatomy
The copulatory bursa, requires skilled dissection and cannot be examined without killing. It is not illustrated here, see Reid (1996) for details and diagrams.
L. fabalis; half length of jelly gland, not reaching capsule gland.
L. obtusata; extends full length of jelly gland to capsule gland.
The ovipositor can be observed on live females restrained as described in Smith (2012 & 2016). Goodwin & Fish (1977) stated that, in 99% of cases examined by them in Wales, its colour is “to varying degrees black pigmented” on L. obtusata 14Dof flic.kr/p/RfXfWn , while L. fabalis “lacks pigmentation” 15Dof flic.kr/p/RfXfGp , but this was not my experience in Wales 16Dof flic.kr/p/Q2j6nF , and Reid (1996) also found it not always so . The colour is often lost on preserved specimens as the surface frequently sloughs off.
To identify females without dissection, the shell forms and shore zone of each species on the same shore need to be ascertained by examination of some penes on males. This information can also be used to find if local ovipositor colours conform with Goodwin and Fish (1977).
7. Differences in shell morphology and colour
No single shell feature is diagnostic in all situations, and some features intergrade. When several features are considered in combination with habitat detail, it is often possible to give a probable identification, but, for certainty, confirmation by examination of penes is required. Some shores, such as in Denmark and from Kent to Dorset, have problematic shell forms, so penis examination is especially necessary (Reid, 1996). Correlations, validated by penis examination, of species with shell form and colour on specific shores can be found in published previous studies, but should be used with caution as shell form may vary on different parts of the same shore (Reimchen, 1981) and may change from year to year with variation in the climate. Shells of Arctic and subarctic populations differ from those south of the Lofoten Islands; they are less well known and are excluded from this account except where specifically mentioned; further detail in Reid (1996).
7a. Shell colours of both species are often classified with the terms below; percentages are of specimens in a large collection of a mixture of both species from across Britain (Smith, 1976, in Reid, 1996). Some authors give slightly different interpretations of a colour term for each species, and it can be difficult to define the limits of fusca, brownish olivacea and faintly marked dark reticulata. Colours are best viewed on live specimens in water as the shell and the periostracum, which often contributes to the colour, erode and fade readily on dead shells 17Dof flic.kr/p/R5n3xZ .
Olivacea: 55%. Exterior olive green 18Dof flic.kr/p/RfXeZ2 to olive brown 19Dof flic.kr/p/R5mYDB , interior often purplish to brownish. Usually the commonest colour form, and almost diagnostic, of L. obtusata on shores well sheltered from waves. An algal coating on L. fabalis 20Dof flic.kr/p/R2FMv5 may confuse unless scraped off; true olivacea L. fabalis are extremely rare or absent in Britain, but they do exist in at least two places in Galicia, Spain on Zostera.
Reticulata: 33%. Exterior yellow to brown with darker reticulate 21Dof flic.kr/p/PYwgVE , chequered 22Dof flic.kr/p/R2FKwW or zig zag 23Dof flic.kr/p/Rcvdds pattern; interior varied, can be white 21Dof flic.kr/p/PYwgVE , sometimes tinted pink or violet and sometimes with exterior pattern showing near rim within aperture 24Dof flic.kr/p/R2FJeL . Usually the commonest colour form of both species on shores exposed to wave action.
Citrina: 10%. Exterior yellow 25Dof flic.kr/p/Rcvc5A grading to white; interior white 26Dof flic.kr/p/R3LhL5 . Usually the commonest colour of L. fabalis on shores well sheltered from waves, but not diagnostic as it also occurs on L. obtusata 27Dof flic.kr/p/R3Lhcj on the same shores.
Fusca: dark brown to black on L. obtusata 19Dof fig.5 flic.kr/p/R5mYDB ; dark chestnut brown on L. fabalis (Reid, 1996). Rare, generally less than 1%, but 100% of sample of 14 L. obtusata s.l. from a site on brackish Isefjord, Denmark (Rasmussen, 1973). A sample of 8 thin shelled, dwarf, L. fabalis from exposed cliff near Aberdeen all appear to be form fusca, but the specimens, at 28Dof flic.kr/p/PZBeay , were photographed 45 years after death, so the colour may have altered.
Each of following less than 1%.
Aurantia: orange. 29Dof flic.kr/p/PZBdiU
Rubens: red or brick red. 30Dof flic.kr/p/QGHGD9
Inversicolor: two broad dark bands. 31Dof flic.kr/p/PZBccW
Zonata: single pale band around periphery (“equator”) of bodywhorl. 33Dof flic.kr/p/R6s4Fe (right specimen).
Alternata: two pale bands. 33Dof flic.kr/p/R6s4Fe (left specimen).
On some shores, juvenile L. fabalis up to 4mm diameter are pure white resembling the 3.5mm diameter spiral tubes of spirorbid worms living on the fronds of Fucus serratus frequented by L. fabalis. With later growth of another colour, the juvenile shell is preserved as a pure white early spire 23Dof flic.kr/p/Rcvdds & 34Dof flic.kr/p/QGHFW7 . Care is required as the spire of L. obtusata is often eroded to dingy whitish, though careful examination will often show traces of the eroded colour 35Dof flic.kr/p/Rdzcqb . And populations of L. fabalis lacking white spirorbid like spires can weather to dingy white with traces of colour in the same way as L. obtusata 36Dof flic.kr/p/R6s3wF .
7b. Shell sizes
On most shores, L. obtusata has a larger mean height than L. fabalis, but there is usually a large overlap in sizes of the largest L. fabalis specimens and young specimens of L. obtusata 37Dof flic.kr/p/R3LbTo . The mean size of both species varies greatly with local conditions 44Dof flic.kr/p/Q3o4dM .
7c. Shell spires
In Britain and most of Atlantic Europe, generally, but not consistently enough for reliable differentiation, L. fabalis has a flat or very low spire 36Dof flic.kr/p/R6s3wF and L. obtusata a low 38Dof flic.kr/p/R6s1kr or slightly raised domed spire 18Dof flic.kr/p/RfXeZ2 , often with an angled shoulder on the body whorl giving a squared or barrel like profile 38Dof flic.kr/p/R6s1kr . Juveniles of both species usually have flatter spires than adults 31Dof flic.kr/p/PZBccW , 32Dof flic.kr/p/R6s4KH , 39Dof flic.kr/p/QGHEsW & 50Dof flic.kr/p/R3L2LW .
Occasional specimens 40Dof flic.kr/p/R6s1gD or local populations have a protruding pointed spire, especially in sheltered brackish conditions, e.g. L. obtusata var. aestuarii in the tidal River Deben, Suffolk, in Jeffreys (1869) fig. 8, plate CI at archive.org/stream/britishconcholog05jeffr#page/n489/mode... (now scarce or extinct at that locality). On Arctic and subarctic shores north of the Lofoten Islands, and in Greenland and most of Iceland, many of both species have protruding pointed spires. Only L. obtusata occurs in Atlantic North America; it often has a developed spire in northern Maine and Canada.
7d. Shell apertures
Aperture size is affected by the physical and biological environment, and juveniles have a less expanded body whorl and aperture. The shell walls thicken with maturity in both species, constricting the aperture internally, generally more markedly in L. fabalis, but varying in degree with environment. The most useful measure of shell thickness is that of the lip at the base of the columella (C) divided by the length of the aperture (LA) 41Dof flic.kr/p/R6rZHz . In a sample of 59 adult shells, including less frequent extreme forms, C/LA was usually less than 0.29 on L. obtusata and greater than 0.29 on L. fabalis, but only with a 75% accuracy (Reid, 1996). Accuracy was greater if extreme forms were excluded. Juveniles of the two species are often very similar with thin shell walls, a sharp, fragile aperture rim and an unconstricted opening, and often the anterior (base of aperture) is drawn out into a moderate spout. As small juveniles lack penes, identification cannot always be verified. Association with verified adults, and shell size and colour, if on a shore where there is a large interspecific difference in these features, are usually the best guide 31Dof flic.kr/p/PZBccW & 32Dof flic.kr/p/R6s4KH .
Graham (1988) states that “a notch where the outer lip and last whorl meet” is indicative of L. obtusata, but its presence varies with age/spire development and it is not a reliable predictor of species (D. Reid, in litt.) 42Dof flic.kr/p/QGHDiw & 43Dof flic.kr/p/PZB3Fs .
8. Phenotypes of different wave exposures
On British rocky shores that have full marine salinity and fucoid algal growth, shell sizes and dominant colours of L. obtusata and L. fabalis vary with the degree of wave exposure. Image 44Dof flic.kr/p/Q3o4dM and its caption summarise approximately the most frequent correlations.
8 a. Sheltered shores, not estuarine, scale 8, 7 or 6, are the best for initial experience of differentiating the two species, providing the tidal range is sufficient to clearly separate and define a zone of Ascophyllum nodosum on the middle/upper shore and a zone of Fucus serratus on the lower shore (low spring tide required to expose) 45Dof flic.kr/p/QGHCb1 . Olivacea L. obtusata at its largest will probably be on the Ascophyllum, and easily differentiated from citrina L. fabalis at its smallest with thick aperture walls on the Fucus serratus 46Dof flic.kr/p/R6rYa4 . Unbiased samples can be obtained by shaking plants into a bucket. Juveniles of both species may be yellow and have a similar shape with an anterior spout 37Dof flic.kr/p/R3LbTo , but juveniles from Ascophyllum with adult L. obtusata will likely be that species and be as large as adult L. fabalis, and far larger than tiny juveniles of L. fabalis. If findings are as described, the identifications will almost certainly be correct, but examining the penes will add to experience. Sites with phenotypes as described can be found on the narrow inner portion of the Menai Straits between the two bridges and probably extending to Bangor and Y Felinheli; aerial photograph at data.nbn.org.uk/imt/?mode=SPECIES&species=NHMSYS00210... .
L. fabalis does not eat Fucus serratus, but uses the flat fronds as suitable feeding platforms for its diet of micro epiphytes and detritus. It is sometimes absent in sheltered situations, despite the presence of F. serratus, if excess sediment coats the plants, e.g. possible cause of absence in upper reaches of Severn Estuary (Williams, 1994). Where turbidity and sediment in estuaries 47Dof flic.kr/p/Rh3x7n or impact of sand laden currents 48Dof flic.kr/p/Rh3wNX prevent the growth of intertidal fucoid algae, both mollusc species are usually absent or very scarce.
8 b. Semi exposed shores, Ballantine scale 4, will provide experience at the opposite end of the phenotype sequence. Ascophyllum, a favoured food of L. obtusata, will be absent or present as a few scattered stunted plants, and L. obtusata will likewise be absent or present as small specimens on scant Fucus vesiculosus or other algae. Fucus serratus is still usually common at scale 4 and L. fabalis usually achieves its largest size here, sometimes equalling or exceeding any L. obtusata present. Differentiating the species can be very difficult on such shores because their sizes are often similar, the predominant colour form of both species is reticulata, and both tend towards larger apertures with thinner shells. Examination of penes is very necessary. An aerial photograph of this sort of shore open to a fetch across the Irish Sea of 100km to 175km, with a wide wave cut platform and plenty of F. serratus but no observed Ascophyllum, is at data.nbn.org.uk/imt/?mode=SPECIES&species=NHMSYS00210...! 0951H+G!0851H+G Large, reticulate L. fabalis were common here (penes checked). The aperture lip of adults was strongly thickened and the opening correspondingly narrowed. The interior was white 21Dof flic.kr/p/PYwgVE , sometimes tinted pink or violet and the exterior pattern often showed within the rim 49Dof flic.kr/p/QGHAn1 . No L. obtusata were found because they were absent or not detected among very similar L. fabalis. A thin shelled, wide apertured, citrina juvenile L. fabalis was initially mistaken for L. obtusata, but the white spirorbid like initial whorls and its presence among adult L. fabalis strongly suggested it was L. fabalis 50Dof flic.kr/p/R3L2LW .
8 c. Exposed shores, Ballantine scale 3, 2 & 1, usually lack either species, but L. fabalis may occur in dwarf form on scale 3 shores 51Dof flic.kr/p/QGHA9q & 28Dof flic.kr/p/PZBeay if sufficiently moist micro habitats exist with secure refuges for dwarfs to retreat into from violent waves. On some Scottish exposed shores with frequent splash and spray, it “may be found further up on the shore on other fucoids where there is adequate protection from desiccation” (McKay & Smith, 1979). It even sometimes occurs at mean high water neap level on Fucus spiralis (Sacchi, 1969 in Reid, 1996). It also occurs in exposed positions on Mastocarpus stellatus in Galicia, Spain; Palmeria palmata at Grindvik, Iceland; Devaleraea stellatus on Achill Island, Ireland; and on red encrusting algae and bare rock in northern Norway (Reid, 1996). On exposed shores, usually, the aperture is larger and the shell thinner than in other situations, and may resemble L. obtusata s.s. from moderately exposed shores.
In a sample of eight from a scale 3 site near Aberdeen, aerial image at data.nbn.org.uk/imt/?mode=SPECIES&species=NBNSYS00001... , all were fusca dwarfs, height mode 4.2mm, max. 5.5mm, with large apertures and thin shells apart from the wide columellar lip 51Dof flic.kr/p/QGHA9q . The spires varied greatly, some, like many L. fabalis, almost flat , others with a large bulging penultimate whorl well proud of the aperture that resembles many L. obtusata s.s..
9. Collecting and reliable recording, suggestions.
# Disregard stranded shells; their anatomy cannot be verified, they lack helpful habitat detail, the periostracum containing much colour may be eroded, and the shell may be bleached and weathered 17Dof flic.kr/p/R5n3xZ .
# If possible, start with a sheltered shore and then a semi exposed shore to gain experience.
# An individual specimen may have atypical features 51Dof flic.kr/p/QGHA9q or be deformed 52Dof flic.kr/p/Q3nXct , so examine several; for inspection of penes, eight mature ones give a 99% chance of both sexes if there is a 50:50 ratio.
# To obtain all colours and sizes, collect the sample randomly, e.g. by shaking plants into a bucket. If only the largest are selected by eye, the sample may be biased to females as they are larger than males on average, especially L. fabalis on sheltered shores.
# Take separate clearly labelled samples from discrete habitats e.g. Ascophyllum and F. serratus zones. Be aware that some shores have more than one wave exposure; see Reimchen (1981), link in references.
# Transport specimens in sealed plastic boxes dampened, but not awash, with seawater.
Store in a fridge at about 6ºC and examine as soon as possible; L. obtusata s.s. exposes itself more readily when damp rather than when submerged.
# Check any shell based identification with examination of some penes.
# When submitting records to a recording scheme, make sure to include that the penes were examined so that at a future date your record can be separated from the mass of less reliable records. Check with the scheme organizer that the detail will be included
when entered. Records submitted to the Marine Recorder of the Conchological Society of G.B. & Ireland have anatomical notes included and uploaded as integral parts of the record; contact marine@conchsoc.org . Other detail such as shore exposure and algal zone are also helpful and welcomed. Clear photographs of shell aperture and penes, or preserved specimens, are valued by the Marine Recorder.
10. Acknowledgements
I am indebted to Dr David Reid for his careful examination of this account and for his highly valued advice, but any errors or omissions are my (IFS) responsibility.
I wish to thank Simon Taylor, Marine Recorder for the Conchological Society of G.B. and Ireland, for helpful discussion and the provision of specimens and photographs, and I am grateful to Dr Jan Light for help with literature and specimens.
11. References and web links
Ballantine, W.J. 1961. A biologically-defined exposure scale for comparative description of rocky shores. Field studies 1(3): 1-19. www.field-studies-council.org/resources/field-studies-jou...
Carvalho, J., Sotelo, G., Galindo, J. & Faria, R. 2016. Genetic characterization of flat periwinkles (Littorinidae) from the Iberian Peninsula reveals interspecific hybridization and different degrees of differentiation. Biol. J. Linn. Soc., 118: 503 to 519. onlinelibrary.wiley.com/doi/10.1111/bij.12762/abstract
Goodwin, B.J & Fish, J.D. 1977. Inter- and intraspecific variation in Littorina obtusata and L. mariae (Gastropoda, Prosobranchia). J. Moll. Stud. 43: 241 to 254. Extract at mollus.oxfordjournals.org/content/43/3/241.extract
Graham, A. 1988. Molluscs: prosobranch and pyramidellid gastropods: keys and notes for the identification of the species. Brill & Backhuys, for Linn. Soc. Lond. & Estuarine and Brackish-water Sciences Assoc. Synopses of the British Fauna (New Series) no.2. Edition 2 (662pages). Leiden.
Hayward, P.J. & Ryland, J.S. (eds.) 1990. The marine fauna of the British Isles and North-West Europe. Volume 2. Clarendon Press, Oxford.
Hayward, P.J. & Ryland, J.S. (eds.) 1995 and reprints to 2009. Handbook of the marine fauna of North-West Europe. Oxford University Press, Oxford.
Jeffreys, J.G. 1865. British conchology. vol.3. London, van Voorst. (L. fabalis included in L. obtusata as var. fabalis.)
archive.org/stream/britishconcholog03jeffr#page/356/mode/1up
Jeffreys, J.G. 1869.British conchology. vol.5. London, van Voorst. Plate ci, fig.8 of Littorina obtusata var. aestuarii (As Littorina aestuarii) from Shottisham Creek, R. Deben, near Felixstowe, Suffolk. Abundant there between tide marks. Well developed spire. (Now extinct or nearly so.) archive.org/stream/britishconcholog05jeffr#page/n489/mode...
McKay, D.W. & Smith, S.M. 1979. Marine Mollusca of East Scotland. Royal Scottish Museum, Edinburgh. [An early adoption of the differentiation as L. littoralis (for L. obtusata s.s.) and L. mariae (for L. fabalis) before nomenclature became settled. Maps reliable as all entries made or vetted by the experienced authors.]
Reid, D.G. 1996. Systematics and evolution of Littorina. Ray Society, London.
www.raysociety.org.uk/publications/zoology/the-systematic...
Reimchen, T.E. 1981. Microgeographical variation in Littorina mariae Sacchi & Rastelli and a taxonomic consideration. J. Conch. Lond. 30: 341 to 350.
www.web.uvic.ca/~reimlab/Microgeographical%20Variation-00...
Sacchi, C. F. & Rastelli, M. 1966. Littorina mariae, nov. sp.: les differences morphologiques et ecologiques entre "nains" et "normaux" chez l'espece L. obtusata (L.) (Gastr., Prosobr.) et leur signification adaptive et evolutive. Atti Soc. Ital. Sci. Nat. 105: 351 to 369.
Smith, D.A.S., 1976. Disruptive selection and morph-ratio clines in the polymorphic snail Littorina obtusata (L.) (Gastropoda: Prosobranchia). J. Mollus. Stud. 42: 114 to 135.
Smith, I.F. 2012. Anatomy of marine gastropods without dissection.
Mollusc World 28: 13 to 15. Conch. Soc. GB & Ireland.
Smith, I.F. 2016. Revision of Smith, I.F. 2012. Anatomy of marine gastropods without dissection. flic.kr/s/aHskNP6GoL and pdf version at www.researchgate.net/publication/310467378_Anatomy_of_mar...
Williams, G.A. 1990. The comparative ecology of the flat periwinkles, Littorina obtusata (L.) and L. mariae Sacchi et Rastelli in Field Studies 7: 469 to 482. fsj.field-studies-council.org/browse-by-category/marine-b... (scroll down to 1990) [Fig. 2 has an exposure scale in reverse order of Ballantine scale so 1 = Ballantine 8; and caption error: closed boxes are L. obtusata, open boxes are L. mariae/fabalis.]
Williams, G.A. 1994. Variation in populations of Littorina obtusata and Littorina mariae (Gastropoda) in the Severn Estuary. Biol J Linnean Soc 51: 189 to-198.
onlinelibrary.wiley.com/doi/10.1111/j.1095-8312.1994.tb00...
12. Glossary
3Dof = Image 3 in Flickr album.
allometric (adj.) = of disproportionate growth of a part or parts of an organism as the whole organism grows.
aperture = mouth of gastropod shell; outlet for head and foot.
coll. = in the collection of (named person or institution; compare with leg.).
columella = solid or hollow axial “little column” around which gastropod shell spirals; hidden inside shell, except on final whorl next to lower part of inner lip of aperture where hollow ones may end in an umbilicus or siphonal canal.
columellar (adj.) = of or near central axis of coiled gastropod.
columellar lip = lower (abapical) part of inner lip of aperture.
columellar muscle = large muscle connecting foot/head of gastropod to its shell at the columella.
distal = positioned or facing away from midline of body.
epiphyte = an organism growing on a larger plant for support, but not nutrition.
epizooic (adj.) = of non-parasitic organisms living on surface of animals.
filament = a slender threadlike object or fibre in animal or plant structures.
filament (in Reid, 1996) = glandless tip of penis a.k.a. distal tubule. Often not threadlike.
flagellum = threadlike organ or appendage.
height (of gastropod shells) = distance from apex of spire to base of aperture along/parallel to axis of coiling (see flic.kr/p/R6s1kr ).
in litt. (abbreviation of Latin “in litteris”) = in unpublished correspondence.
leg. (abbreviation of legit) = collected/ found by (compare with coll.)
mamilliform = shaped like a nipple.
mesial = facing towards the midline of the body.
mobile = able to move self, e.g. a horse, or to be moved by an agent, e.g. a wave or a cell phone.
motile = moves self spontaneously without volition; applicable to whole animals or to parts of them such as cells, gametes, penes or cilia.
penes (plural of penis) = male copulatory organs.
phenotype = form of a species resulting from influence of particular environmental factors.
proximal = towards the centre of the body or point of attachment.
sensu lato (abbreviation s.l.) = in the wide sense, possibly an aggregate of more than one species.
sensu stricto (abbreviation s.s.) = in the strict sense, excluding species that have been aggregated or confused with it.
spire = all whorls of a gastropod shell, except the final body whorl. But in this account, “spired shells” are taken to be those with a pointed apex protruding beyond the body whorl rather than the domed spire of some that extends beyond the body whorl.
taxonomic (adj.) = of or relating to taxonomy.
taxonomy = the description, identification, naming, and classification of organisms.
vermiform = shaped like a worm.