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funny farewell quotes for boss
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Site InfoWe are the home of 1295 authors from among our 2601 members. There have been 2834 reviews written about our 3854 stories consisting of 10945 chapters and 29671511 words. A special welcome to our newest member, Ahewi17.
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Secret Hearts, Secret Love by Aglarien
Summary: Elladan and Elrohir have had enough of deception.
Rating: Teenage Audience [Reviews - 0]
Category: FPS > Elladan/Elrohir, FPS, FPS > Elrond/Erestor, FPS > Erestor/Elrond, FPS > Elrohir/Elladan
Characters: Elladan, Elrohir, Elrond, Erestor
Type: None
Warning: AU
Challenge: None
Series: None
Chapter: 1 | Completed: Yes | Words: 673 | Read: 1369
Published: November 04, 2008 | Updated: November 04, 2008
Story notes: Beta: *whistles in the wind*
1. Chapter 1 by Aglarien [Reviews - 0] (673 words)

subset: commoncrawl


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Sold Out
Santoku means "the 3 virtues" (slicing, dicing, mincing) and stands for this knife's high versatility. It is the perfect multi-purpose knife for vegetables, meat and fish. This great universal tool isn't as intimidating as a large chef knife and therefore perfectly suited to initiate you to the great world of Japanese knives.
Handmade Masakage Kiri knives are made of VG10 folded Damascus stainless steel, which guarentees easy use and low maintenance. The knife has been named Kiri (Japanese term for "Mist"), in reference to the blade's matt and foggy finish. Its oval handle is made of lightweighted Magnolia and red Pakka Wood. As with most Masakage knives, the heel of the blade is rounded to allow a more comfortable use of the pinch grip.
All knives from Masakage are handmade by master blacksmiths of Takefu's village and hand sharpened by one of Japan's best knife sharpeners, Takayuki Shibata san. When these knives come out of their box, they have an amazing sharpness and will stay sharp for an extremely long time due to Shibata san's masterful technique. Alambika is extremely happy to import and retail Japanese knives (from Takefu village) in Canada. The whole kitchen knife collection can be seen and handled in our Montreal store.
Be the first to ask a question about this.


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John Surman — Way Back When
(Cuneiform Rune 200, 1969/2005, CD)
by Jon Davis, Published 2006-05-01John Surman (saxes), John Marshall (drums), John Taylor (electric piano), Brian Odgers (bass guitar). For many readers, that list alone puts this on the must-have list. Throw in the recording date of fall 1969, and the mind starts calculating. Think of early Nucleus, think of electric Miles, think of the possibilities. This outstanding recording meets and exceeds those possibilities, being a crystal clear slice of history, when jazz and rock were first starting to come together. And while the sessions were an informal affair between a few musical friends, the result is top-notch, live and spontaneous and grooving as all get-out. Aside from a few test pressings, this material never saw the light of day until the master tapes, thought long lost, were unexpectedly rediscovered in 2003, so this has to be one of the most extraordinary rarities. The sound is amazingly pristine. Surman's playing is masterful, expressive and melodic at times and wildly intense at others, imaginative and fluid. John Marshall is, well, John Marshall, one of England's finest jazz drummers. Odgers is very good here, and while much of his resume is in rock and pop, he was in John McLaughlin's Extrapolation band with Surman. Taylor is on fire at the keyboard, bringing out the peculiar tones of the electric piano and excellent on both his solos and his backing of the others. This is one forgotten gem that we are fortunate to have available, surely one of the best reissues of recent years.
Filed under: New releases, Issue 33, 2005 releases, 1969 recordings
Related artist(s): John Surman
Celebrate 10 Years of Fruits de Mer – As a special celebration for a decade of cool vinyl releases, our friends at Fruits de Mer records have prepared a limited edition reissue of an album by the first band ever to appear on the label: Schizo Fun Addict. The band is known for unusual release strag » Read more
Mega Dodo Presents New Charity Album – Our friends at Mega Dodo have put together a lovely compilation of their artists performing new arrangements of nursery rhymes, and all the profits from sales of the album will benefit Save the Children. It features a number of artists we've covered. » Read more
Phil Miller RIP – Sad word reaches us today of the passing of another of the great musicians of the Canterbury Scene — guitarist Phil Miller. His distinctive sound added greatly to Matching Mole, Hatfield and the North, and National Health, and he also contributed to albums by Caravan, Dave Stewart & Barbara Gaskin, and many others. He was 68. » Read more
Moonjune to Distribute Tony Levin's Back Catalog – It has been announced that Moonjune will now handle distribution for Tony Levin's catalog of releases. These great albums will now be a bit easier to get hold of, so check out the site and see what you're missing. The veteran of King Crimson and Stick Men worked with a host of great players on these albums, and we've reviewed most of them over the course of the years. » Read more
Bandcamp Shines Light on Niches We Like – Bandcamp has developed into one of the best places to discover new music, and even a lot of old music is showing up there. In addition, their staff has been producing periodic articles spotlighting some interesting stylistic areas. On 20 September, they published one called "The New Face of Prog Rock" which bears checking out. » Read more
Hinn Íslenzki Þursaflokkur - Þursabit – This band hails from Iceland, and to get a hint at their sound you might think of a Scandinavian Gryphon mixed with a large dose of Gentle Giant. This was their second album, released in 1979. Hinn... (1994) » Read more


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Pick of the Week
Meet our Guest Editor
Stuart Crewes, Creative Director of Art Week Exeter, selects our Pick of the Week feature
Artist of the Month
Latest news
Artist Commission opportunity at Cothele
Monday 20 November 2017
We are looking to commission an artist or crafts person to create a new work to be integrated within or displayed in conjunction with the Cotehele garland for 2018.
Statement on the closure of alias - the Artist-Led Initiative Advisory Service
Tuesday 24 October 2017
The Artist-Led Initiative Advisory Service (alias) has announced that it has ceased all activities effective from October 2017.
Visual Arts South West is a network creating opportunities for artists, organisations and professionals to develop their practice, share ideas, knowledge & resources, and cultivate relationships. We provide platforms to promote and support the work of the region's visual arts ecology.
We are part of the national Contemporary Visual Arts Network (CVAN) which is dedicated to working together to realise a stronger visual arts for the benefit of artists, arts organisations and audiences, present and future.
Featured profile
The Royal Photographic Society
What's on in the South West
The Vestibules, City Hall, College Green, Bristol, BS1 5TR
Monday 20 November 2017 – Friday 24 November 2017
CAST Film Club: Phil Collins
CAST, 3 Penrose Road, Helston, TR13 8TP
Friday 24 November 2017
Odyssean: Topographies
Hestercombe Gallery, Cheddon Fitzpaine, Taunton, Somerset, TA2 8LG
Saturday 18 November 2017 – Sunday 25 February 2018
Mixing Signals
Kobi & Teal, 57 Catherine Street, Frome, Somerset, BA11 1DA
Wednesday 01 November 2017 – Saturday 23 December 2017
Silk-Screen Printed Pocket Squares
The Holburne Museum, Great Pulteney Street, Bath BA2 4DB
Saturday 09 December 2017
Don Mandarin: An Occult Diary
Centrespace Gallery, Bristol, BS1 1EA
Saturday 25 November 2017 – Wednesday 29 November 2017
Two Dimensions
Harbour House, The Promenade, Kingsbridge, TQ7 1JD
Tuesday 21 November 2017 – Sunday 03 December 2017
5weeks stone carving with Sarah Klopper
Dorset Centre for the Creative Arts Middle Farm way, Poundbury, Dorset DT1 3AR
Tuesday 31 October 2017 – Tuesday 28 November 2017


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Aon Employee Benefits acquires Portus Consulting
Aon Employee Benefits has acquired Portus Consulting, a independent UK-based employee benefits firm, for an undisclosed sum.
Portus specialises in the legal sector and among SMEs with 50 to 750 employees. The acquisition of Portus supports Aon's ambitions to grow its UK employee benefits business both organically and through acquisition. It will join Aon Employee Benefits, which provides consultancy, broking services and technology to 2,500 large and SME corporate clients, covering over 1 million lives in the UK. Terms have been agreed, subject to approval by the Financial Conduct Authority (FCA), and Aon anticipates...
More news
British Friendly introduces Adviser Toolkit
British Friendly has introduced an online Adviser Toolkit to encourage income protection sales.
Zurich launches 'selfie' app to encourage life insurance take-up
Zurich has launched a 'selfie' app called FaceQuote which estimates how much life cover someone might need.
COVER Excellence Awards 2017 - Winners supplement out now
The official supplement of this year's COVER Excellence Awards is available to read now as an eBook.
CIExpert: An analysis of children's critical illness benefits
Child cover within critical illness plans has assumed a far greater importance in recent years, writes CIExpert's Alan Lakey.


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IRVING (AP) – Retail gasoline prices across Texas continue to fall, with the per-gallon cost dropping two cents over the last week.
AAA Texas on Thursday reported the average statewide unleaded price at the pump is now $3.09. The price has fallen by a nickel over the last two weeks.
The statewide average is 16 cents less per gallon than the national average of $3.25.
Of the metro areas in Texas surveyed by AAA, drivers in Dallas are paying the most at $3.12 and drivers in Amarillo continue to pay the least at $2.89.
Retail gas prices nationwide generally moved lower following the end of the summer driving season in September due to plentiful supplies, flat demand and falling crude oil prices.
(© Copyright 2013 The Associated Press. All Rights Reserved. This material may not be published, broadcast, rewritten or redistributed.)
Latest News:
- No. 16 Texas A&M Routs Oklahoma State 72-55 At Tournament
- No. 22 Baylor Beats Wisconsin 70-65
- Judge Permanently Blocks Trump Sanctuary Cities Order
- Texas Longhorns Bowl Eligible For 1st Time Since 2014
- Rare 'Dracula' Poster Sells For $525,800 At Dallas Auction
Top Trending:
- Cowboys Fan Attacked, Beaten After Thanksgiving Day Game
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- PHOTOS: Your Pet Pictures


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Behind the Mask exhibition hits Liverpool
Tony Murphy, BT Buddies Trustee and Fifi, Brain Tumour Survivor
I don't think there is anyone who attended the Liverpool Launch that won't know who Fifi is! What a little superstar!
Fifi had radiotherapy for a brain tumour when she was just 3 years old, her mask is tiny and decorated like a little princess! Fifi, her Mum Mya and her Jamjar (Grandma) made the 2 hour trip to our exhibition and it was so lovely to meet them all!
There were other visitors that made long trips to be in Liverpool for our exhibition launch. Michael, a Glioblastoma Multiforme survivor and contributor to our exhibition brought his Mum, Step Dad and brother along to see his mask and the rest of the exhibition. Thanks to Step Dad for taking a few photos for us!
We would like to thank Gilly for her helping in setting everything up and Claire for going above and beyond by staying on to hand out drinks on launch evening!
The event went well and we hope we reached a few people who may otherwise not have heard about brain tumours.


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More Videos...
Super Fast EVO - Uncontrollable - TX2K12
These 2 Mitsubishi EVOs were crazy fast! I can't believe the silver one was running over 150mph They were kind of like the black Supra in the last video, they were all over the track and looked like they might crash a few times. http://www.tx2k.com Check out http://www.hightechcorvette.com http://www.houstonperformancedriving.com HTC Facebook http://www.facebook.com/hightech.corvette HPD Facebook: http://www.facebook.com/HoustonPerformanceDriving Twitter: http://twitter.com/HTCorvette Fan Page: http://www.facebook.com/HighTechCorvettefans
ILR EVO Dyno Run
ILR EVO IX on a Dyno @ 40 PSI. Was done using a MD Dyno.
Mitsubishi Lancer Evolution IX @ LancerEvoClub.org Verhoeven Tuning Rollenbankdag
Mitsubishi Lancer Evolution IX @ LancerEvoClub.org Verhoeven Tuning Rollenbankdag 19-6-2010
Nissan GTR vs Mitsubishi EVO FQ 400 - Fifth Gear
In this classic Fifth Gear clip these two Japanese super cars are going head to head to see which is best, first is an autotest and the second a 2 lap race! For more fantastic car reviews, shoot-outs and all your favourite Fifth Gear moments, subscribe to our Official Channel - https://www.youtube.com/user/fifthgearuk


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Emergency shelter doubles as a ground sheet. Can be used as a distress signal or for storing gear. Also helps prevent hypothermia. Made from 3-mil polyethylene.
Size: 84" x 36"
Weight: 9 oz.
Size: 84" x 36"
Weight: 9 oz.
Alt Information
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1 - 3 Days
Sleep Gear
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- Bivvies & Blankets
Customer Reviews
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Anion exchange silica monolith for capillary liquid chromatography
- 175 Downloads
- Citations
An anion exchange monolithic silica capillary column was prepared by surface modification of a hybrid monolithic silica capillary column prepared from a mixture of tetramethoxysilane (TMOS) and methyltrimethoxysilane (MTMS). The surface modification was carried out by on-column copolymerization of N-[3-(dimethylamino)propyl]acrylamide methyl chloride-quaternary salt (DMAPAA-Q) with 3-methacryloxypropyl moieties bonded as an anchor to the silica surface to form a strong anion exchange stationary phase. The columns were examined for their performance in liquid chromatography (LC) and capillary electrochromatography (CEC) separations of common anions. The ions were separated using 50 mM phosphate buffer at pH 6.6. Evaluation by LC produced an average of 30,000 theoretical plates (33 cm column length) for the inorganic anions and nucleotides. Evaluation by CEC, using the same buffer, produced enhanced chromatographic performance of up to ca. 90,000 theoretical plates and a theoretical plate height of ca. 4 μm. Although reduced efficiency was observed for inorganic anions that were retained a long time, the results of this study highlight the potential utility of the DMAPAA-Q stationary phase for anion separations.
KeywordsAnion exchange Silica monolith Micro liquid chromatography Capillary electrochromatography Inorganic anions
Capillary electrochromatography
Methyl trimethoxysilane
N-[3-(dimethylamino)propyl]acrylamide methyl chloride-quaternary salt
- 1.Paull B, Nesterenko PN (2005) TrAC 24:295–303Google Scholar
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- 4.Zakaria P, Hutchinson JP, Avdalovic N, Liu Y, Haddad PR (2005) Anal Chem 77:417–423CrossRefGoogle Scholar
- 5.Riordain CO, Gillespie E, Connolly D, Nesterenko PN, Paull B (2007) J Chromatogr A 1142:185–193CrossRefGoogle Scholar
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- 7.Hatsis P, Lucy CA (2003) Anal Chem 75:995–1001CrossRefGoogle Scholar
- 8.Ikegami T, Ichimaru J, Kajiwara W, Nagasawa N, Hosoya K, Tanaka N (2007) Anal Sci 23:1–6CrossRefGoogle Scholar
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EA allegedly considering two ways of re-introducing microtransactions into Star Wars Battlefront 2; Details
Nov 17, 2017 | 18 Comments
4J Studios has revealed the latest batch of fixes for the PlayStation 3 version of Minecraft. Included in patch 1.03 are fixes for a number of game crashing issues which 4J already posted. The developer hopes the latest update will be available before next week, but that all depends on how it does in Sony's test. In the meantime, check out what you can expect in the upcoming update.
Change log for Minecraft: PS3 Edition Patch 1.03
The PS3 version of Minecraft was released in mid-December 2013. PlayStation 4 and Vita versions are also in the works, though neither has a release date other than some time this year.


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Known as America's stonehenge, where did the Georgia Guidestones come from, who put them there and why?
The Georgia Guidestones is a granite monument erected in 1980 in Elbert County, Georgia, in the U.S. A set of 10 guidelines is inscribed on the structure in eight modern languages, and a shorter message is inscribed at the top of the structure in four ancient language scripts: Babylonian, Classical Greek, Sanskrit, and Egyptian hieroglyphs.
Dr. J. Michael Bennett is the primary researcher and the associate producer behind a new documentary film:"Dark Clouds Over Elberton: The True Story of the Georgia Guidestones."
In this interview James Corbett talks to Michael about the mystery surrounding the Guidestones.
"Combining new interviews with some of the key people involved in the guidestones construction with never-before-seen evidence and good old gumshoe detective work, the documentary reveals the identity of the guidestones' pseudonymous creator, "R.C. Christian," and in the process uncovers dark connections to the racist, Malthusian pseudoscience of eugenics."
Below is the trailer for the new documentary by Adullam Films
Carol Adl
Latest posts by Carol Adl (see all)
- Sylvester Stallone Denies Claims He Sexually Assaulted Teenage Fan In 1986 - November 17, 2017
- Israel Will 'Act Alone' Against Iran In Syria Warns Netanyahu - November 16, 2017
- London's Old Vic Theatre Reveals 20 Staff Allegations Against Kevin Spacey - November 16, 2017


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Guest House idea: I love how the bed is above the living space. Great idea for a small space!
White house, black shutters, red door, partial wrap-around porch.
The decor live serves you all your interior home and office decoration furniture ! Even if you are looking to party decoration furniture and accessories.
Living room seating HGTV Dream Home 2015 - Wood Beams on Ceiling


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- More
[Hartmann Schedel. Liber chronicarum. Nuremberg: Anton Koberger for Sebald Schreyer and Sebastian Kammermeister,...Click the image to load the highest resolution version.
First Edition of the Nuremberg Chronicle, the Greatest Illustrated Book of the Fifteenth Century[Hartmann Schedel. Liber chronicarum. Nuremberg: Anton Koberger for Sebald Schreyer and Sebastian Kammermeister, 12 July 1493].
First edition of the Nuremberg Chronicle, preceding the German language edition (published 23 December 1493) by just over five months. Large folio (17.125 x 11.5 inches; 436 x 292 mm.). 325 (of 328) leaves ([1, title], [19, index], CCXCIX, [1], [5]), lacking only three blank leaves (the two final blanks, as well as the blank leaf following the five unnumbered leaves). Preliminary leaves [6] and [7] are mounted on stubs and supplied from another copy, leaf [7] bound following leaf [20]; a few additional leaves possibly supplied from other copies. This copy contains the three numbered leaves CCLVIIII-CCLXI (259-261), blank except for headlines (often lacking), and the five additional unnumbered leaves containing the description of Poland, De Sarmacia regione Europe, and laudatory verse on Maximilian, here bound at the end. Foliation corrected (with paint or a stamp) on leaves CCXXIX (229) and CCXLVII (247). Gothic type, sixty-four lines plus headline. Table and parts of the text in double columns. Woodcut title and 1,809 woodcut illustrations, of which 1,164 are repeats, from 645 blocks (Sydney Cockerell's count in Some German Woodcuts of the Fifteenth Century (Hammersmith: Kelmscott Press, 1897), pages 35-36), by Michael Wolgemut, Wilhelm Pleydenwurff, and their workshop, including the young Albrecht Dürer, including double-page maps of the World (Shirley 19) and of Europe by Hieronymus Münzer after Nicolas Khrypffs, twenty-nine double-page city views, and eight full-page woodcuts. Most of the woodcuts have been colored by hand at a later date. The double-page map of Europe and the double-page view of "Nissen" in the Sarmatian supplement are uncolored. Two- and three-line printed lombard initials. Two- to fourteen-line capital spaces; several initials in the index and a few initials in the text have been supplied or supplied in outline in red; capital strokes, paragraph marks, and underlining supplied in red.
The portrait of Pope Joan ("Joannes septimus," verso of leaf CLXIX (169)), usually missing, is present and unmutilated. The verso of leaf CCLVII (257) is in the later, corrected state, with the uppercase As present, and the recto of leaf CCLVIII (258) is in the first, uncorrected state, with the uppercase As omitted (presumably Koberger ran short and left spaces for the As in early copes of this sheet, later filled in with a new supply). The colophon at the end of the Sixth Age of the World, on CCLVIII (258) verso, is dated 23 May 1493 ("x. kal[end]as Junias"); the colophon at the end of the Seventh and Last Ages of the World, on CCLXVI (266) recto, is dated 4 June 1493; and the final colophon on the verso of the map of Europe, [CCC] ([300]) verso, is dated 12 July 1493. The first two colophons identify only the author; the final colophon identifies the patrons, the printer, and the artists.
Expertly restored in early blindstamped calf over thick bevelled wooden boards (binding measurements: 18 x 12.375 inches; 457 x 315 mm.). Neatly rebacked, retaining the original spine and two leather pull-straps across the spine. Two metal catch plates on the front cover renewed, metal corner plates on both covers renewed, remains of straps on rear cover (one renewed). Later vellum endleaves from a manuscript choirbook. Boards a bit bowed, some minor worming to covers. The title, which has been backed, is darkened, with a few smudges or stains and a few marginal wormholes; the top blank portion (two to three inches) of the title has been torn away. There is a dampstain in the lower gutter margin throughout much of the book, very faint at the beginning, heavier toward the end, with occasional traces of mold, which, combined with some worming, has necessitated paper repairs to several leaves (neatly accomplished). Additional repairs to the upper and outer margins and corners of preliminary leaf [6]; repaired tear to preliminary leaf [7], affecting four lines of text; repair to leaf XXI (21), affecting a few letters in the last two lines on the recto; repaired tear to the lower margin of leaf CXIV (114); repairs to the lower margin of leaves CLXXXIV (184), CLXXXV (185), CXCVIII (198), and CCLIII (253), just entering the woodcuts; a few additional leaves with neat marginal repairs or strengthening. Several tiny puncture holes in the woodcuts on leaves II, III, and IV (2, 3, and 4), expertly and almost invisibly filled. Two tiny wormholes in the text from leaf CCLXVIII (268) to the end, sometimes affecting a letter or two. Occasional browning; some smudging of the coloring and offsetting or bleeding onto adjacent leaves; a few additional small stains or splatters. Early ink ownership inscription on the title, the following leaf, the first leaf of text, and on the verso of the final colophon leaf, all inked out; a few scattered early ink marginalia and corrections. The double-page map of Europe on leaves CCXCIX (299) verso and [CCC] ([300)] recto has several cities in Germany and Switzerland added in ink in an early hand. Overall, a sturdy, handsome copy.
"The text by Schedel records the history of the world from the Creation to 1493 and-after allowing six blank pages for the purchaser to fill in a brief account of his own time-concludes with the apocalyptic vision of St. John and the Day of Judgment. In its reach the work covered all the aspects of history: biblical, classical and traditional. It is the illustrations on which the fame of the volume rests. There are 1,809 woodcuts printed from 645 different blocks. They picture the major events of the Old and New Testaments, episodes in the lives of many saints, portraits of prophets, kings, popes, heroes, and great men of all centuries, freaks of nature, and panoramic views of cities. An economic use of the blocks resulted in certain curious anomalies: Nebuchadnezzar appeared later as several German emperors, and the town of Mainz does service also for Naples. Nuremberg artists Michael Wolgemut and Wilhelm Pleydenwurff were responsible for the production of the book...The wood blocks were designed by the two masters and their assistants, including the young Albrecht Dürer, who was [godson to Koberger and] apprenticed to Wolgemut at the time. The printing was carried out under the supervision of the great scholar-printer Anton Koberger, whose illustrated books were famous throughout Europe" (Legacies of Genius: A Celebration of Philadelphia Libraries 5).
On the verso of leaf CCXC (290) "is a passage which has been used as an authority for the [unwarranted] claim that Martin Behaim of Nuremberg, in a voyage made by him with James Camus, or Jacob Cam, under orders from the King of Portugal, went to the Southern Sea near the coast of Africa, and, after crossing the equator, reached the New World and discovered new islands. This passage is not in the German edition, and an examination of the original manuscript of the work, which is still preserved in Nuremberg, has shown it to be an interpolation, written in a different hand" (Church 7; see also Harrisse 13 and Sabin 77523).
BMC II, page 437. Bodleian S-108. Fairfax Murray, German, 394. Goff S-307. GW M40784. Hain-Copinger 14508*. Harvard/Walsh 726-732. ISTC is00307000. Polain 3469. Proctor 2084*. Schreiber 5203. See also Adrian Wilson, The Making of the Nuremberg Chronicle (Amsterdam: 1976).
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Millions of doses of yellow fever vaccine will be handed out to curb the spread of the disease in Sudan
Over a million people in Sudan will be offered immunisations in the next month to battle what experts believe is the worst outbreak of yellow fever the world has seen for 20 years. Since the immunisation programme began on November 21 about 2.2 million people have been vaccinated against the disease, which can cause jaundice and kidney failure. More than 800 suspected cases of the disease have been reported since October, prompting a mass vaccination campaign.
With another 2.2 million doses of the vaccine expected to arrive soon, the World Health Organization's Anshu Banerjee said he was confident the immunisation programme would slow the spread down. Neighbouring countries like Cameroon have also reported cases of the disease but many of these nations include yellow fever vaccine in childhood immunisation programmes.
"There's been no such vaccine campaign in Sudan for decades," said Dr Banerjee. "It's because of priorities. Sudan recently identified itself as a high-risk country for a yellow fever outbreak and had conditional approval to begin a vaccination campaign. Unfortunately the outbreak hit before this could happen.This is the worst outbreak we have seen globally in 20 years or more." Visitors to all parts of Africa are recommended to take out appropriate
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18 November 2017
Owning a dog could be the key to reducing the risk of heart attacks and other potentially terminal diseases, according to a major new study.
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Women diagnosed with untreatable breast cancer are being given fresh hope, with the approval of two new "breakthrough" medicines for widespread use across England's health service.
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A diet of fruit, veg and coffee could be the latest step in reducing the risk of heart failure, scientists say.
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Obese people are to be given fitbit-style wristbands to reduce the number of Type 2 diabetes diagnoses.
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A young man who had both his legs amputated after being electrocuted is not only walking again, but is about to set sail on a global adventure.
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Women who have been inoculated against human papillomavirus (HPV) can attend 75% fewer cervical screenings, a new study has found.
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Britons are afforded less face-to-face time with their GP than dozens of other developed nations, a new study has found.
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A new appetite suppressant developed from concepts relating to the Atkins and Dukan diets could be the latest step in tackling obesity, scientists claim.
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text: I've had a couple of bowls recently that were finished drying, and since the rims were nearest the pith (but didn't include the pith, of course) there were the usual peaks in the rims. One of them, I needed to take off a half-inch+, and the wall thickness was a half-inch, so I wasn't sure the best way to take that off. Actually ended up scoring a line with a parting tool, and then using a little saw-knife and then thin-kerf saw (lathe off) to cut through it. Is there a better, more kind-to-the-hands way to do this?


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TVs, tablets, smartphones, and other electronic gadgets can affect how easily children get to sleep. Try to keep yo… https://t.co/ImR36CLjuD
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Read about the symptoms of narcolepsy, including excessive daytime sleepiness, sleep attacks, cataplexy and sleep paralysis.
Not everyone with narcolepsy experiences the same symptoms. Some experience them regularly, while others are less frequently affected.
Symptoms may develop slowly over a number of years, or suddenly over the course of a few weeks.
Narcolepsy is usually a long-term (chronic) condition, although some of the symptoms may improve as you get older.
You should make an appointment to see your GP if you think you may have narcolepsy so they can determine what's causing your symptoms. If necessary, you'll be referred to a sleep disorder specialist, who can confirm the diagnosis.
Read more about diagnosing narcolepsy.
In most cases, excessive daytime sleepiness is the first sign of narcolepsy. This can have a significant impact on everyday life.
Feeling drowsy throughout the day and struggling to stay awake makes it difficult to concentrate at work or school. People with narcolepsy may be misjudged as being lazy or rude.
Sleep attacks – falling asleep suddenly and without warning – are also common in people with narcolepsy. They may occur at any time.
The length of time a sleep attack lasts will vary from person to person. Some people will only have "microsleeps" lasting a few seconds, whereas others may fall asleep for several minutes.
If narcolepsy isn't well controlled, sleep attacks may happen several times a day.
Most people who have narcolepsy also experience cataplexy – sudden, temporary muscle weakness or loss of muscular control.
Typical symptoms are:
Cataplexy attacks are usually triggered by an emotion, such as excitement, laughter, anger or surprise. Attacks can last from a few seconds to several minutes.
Some people with narcolepsy have cataplexy attacks once or twice a year, while others experience them several times a day. In an attempt to avoid attacks, some people may become emotionally withdrawn and socially isolated.
Some people with narcolepsy experience episodes of sleep paralysis. This is a temporary inability to move or speak that occurs when waking up or falling asleep.
The episodes can last from a few seconds to several minutes. Although sleep paralysis doesn't cause any harm, being unable to move can be frightening.
As well as the symptoms described above, narcolepsy can cause a number of other symptoms, including:
Speak to your GP if you have narcolepsy and it's making you feel low or depressed.
Your GP can advise you about how to minimise the effect narcolepsy has on your daily life. They can also put you in touch with a narcolepsy organisation or support group, such as Narcolepsy UK.


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The effect of arbuscular mycorrhizal fungi (AMF) on micropropagated banana plantlets was evaluated during the acclimatization period. Plants were inoculated with Acaulospora scrobiculata, Glomus clarum or Glomus etunicatum. After cultivation in a greenhouse for 3 months, height, leaf area, fresh weight and dry matter of root and shoots, level of AMF… (More)
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Microbiological analyses of chicken eggs in Recife and Salvador have shown a high occurrence of Salmonella in the egg shells and yolks. Likewise, the occurrence of Salmonella plus coagulase-positive staphylococci in Coalho cheese reached alarming levels. The data revealed a significant risk of infections and intoxications from consuming these foods in the… (More)
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Selective pressures that occur during long-distance migration can influence morphological traits across a range of taxa. In flying insects, selection should favour individuals that have wing morphologies that increase energy efficiency and survival. In monarch butterflies, differences in wing morphology between migratory and resident populations suggest that migratory populations have undergone selection for larger (as measured by length and area) and more elongated (as measured by roundness and aspect ratio) forewings. However, selection on wing morphology may also occur within migratory populations, particularly if individuals or populations consistently migrate different distances.
Using 613 monarch butterflies that were collected on the Mexican wintering grounds between 1976 – 2014, we tested whether monarch wing traits were associated with migratory distance from their natal areas in eastern North America (migration range: 774–4430 km), as inferred by stable-hydrogen (δ 2H) and -carbon (δ 13C) isotopic measurements. Monarchs that migrated farther distances to reach their overwintering sites tended to have longer and larger wings, suggesting positive selective pressure during migration on wing length and area. There was no relationship between migration distances and either roundness or aspect ratio.
Our results provide correlative evidence that the migratory period may act as a selective episode on monarch butterfly wing morphology, although selection during other portions of the annual cycle, as well as extensive mixing of individuals from various natal locations on the breeding grounds, likely counteracts directional selection of migration on morphology.
The online version of this article (doi:10.1186/s40462-017-0098-9) contains supplementary material, which is available to authorized users.
The selective pressures associated with long-distance migration can influence morphology in a variety of taxa, including birds, insects and fish [1–4]. Because migration is energetically expensive [3] and high mortality is predicted during the migratory phase of the annual cycle [5, 6], selection should favour individuals that have morphological advantages that increase efficiency and hence migratory success. For example, populations of birds that migrate longer distances have longer and more pointed wings than sedentary populations or populations that migrate shorter distances [7–11]. Long and thin wings with elongated tips improve aerodynamics and reduce drag during gliding flight [12, 13]. The same selection pressures should operate on migratory insects [4, 14], but examining this hypothesis is hampered by the challenges associated with assigning insect origins and tracking long-distance movements.
One way to estimate migratory distance of insects is through the use of endogenous markers such as stable isotopes [15, 16]. Stable isotope relative abundances (e.g. δ 2H, δ 15N and δ 13C) are translated up the food chain [15, 17] and, if they vary in a spatially predicable manner (termed isoscapes; [18]), reflect the geographic origin of where tissues are grown in herbivores and higher order consumers [15]. Since stable isotopes in the chitin of wing tissue of adult flying insects is metabolically inert after formation [15, 19, 20], stable isotope measurements can be used to identify the geographic natal origin of adults regardless of how far they have flown from the natal site [21]. Stable isotope measurements have been used successfully to determine natal origin of migratory insects such as butterflies [22–24] and dragonflies [21].
Monarch butterflies (Danaus plexippus) in eastern North America are known for their fall migration up to 5000 km from their northernmost breeding range in Canada to overwintering sites in central Mexico [25]. Migratory monarchs are known to have longer wings compared to resident populations [26–28]. Individuals with more elongated wings tended to be seen earlier in migration than those with short wings, suggesting that elongated wings increase the speed of migration [29]. At overwintering grounds in Mexico [30] and the west coast of the U.S. [31], there is a negative correlation between hydrogen isotope values (δ 2H) and wing size, suggesting that individuals with larger wings derived from more northern locations and so migrate longer distances. These studies were based on single-year sampling, and recent work showed that patterns of monarch natal origin can vary annually [32]. Furthermore, both studies used δ 2H values as a proxy for migration distance, which, in some cases, can introduce a significant source of variation because δ 2H values do not necessarily correlate directly with distance travelled. At the same time, larger adult wing sizes may arise when larvae develop at lower mean temperatures at northern latitudes [33] and this parameter must be controlled for in statistical analyses.
Here we examined the hypothesis that migration distance acts as a selective episode on monarch wing morphology because of the energetic costs of longer flights. To examine this hypothesis, we estimated migration distance of overwintering monarchs in Mexico sampled over five decades by estimating natal origin from two stable isotopes (δ 13C and δ 2H) in their wing tissue [22, 34–36]. Given that last generation monarchs in eastern North America migrate directly to a few spatially clustered locations in central Mexico [37], fall migration distance and ambient temperature experienced during larval development can be estimated as a direct function of natal origin in eastern North America. We considered that individuals sampled in Mexico to be representative of those that survived the migratory journey. Thus, if migration distance influenced wing morphology by selecting for individuals with more efficient long-distance flight [26], after controlling for temperature we predicted that fall migration distance would be positively related to i) forewing size (as measured by length and area; Fig. 1) and ii) the degree of elongation in the forewing (as measured by roundness and aspect ratio; Fig. 1) in monarchs sampled on their overwintering grounds.
Monarch butterflies (Danaus plexippus) (N=613) from the eastern North American migratory population were collected from the Sierra Chincua Sanctuary in Mexico during the overwintering period (November - February) from 1976–2015; samples from 1988 did not have any colony location recorded (Additional file 1: Table S1). Forewings were detached from body using forceps and scanned on an HP PSC 750 flatbed scanner at 300 dots per inch [38]. ImageJ software [39] was used to take wing morphology measurements (length, width, area) of forewings images following Altizer and Davis [26]. Length was defined as the point of attachment at the thorax to the distal tip of the wing, width was defined as the longest line drawn perpendicular to the length axis, and area was defined as the total surface area of the forewing (Fig. 1). These measurements were then used to calculate wing roundness (4 × [area/(π × length)2]; [38]) and aspect ratio (length/width; [26]). Our measurements of roundness reduced the skew of perimeter measurements that occurred from variance due to wing wear [38] and resulted in data that were normally distributed in order to meet the assumptions of linear models. Individuals with wings that were frayed or damaged that would have resulted in inaccurate forewing length measurements were excluded from our analyses (N=91). The distance measurements using the centroid method (see below) did not vary between the butterflies included and those excluded (t=−1.19, p=0.24). However, in a pattern opposite to our expectations, butterflies with worn wings (mean distance: 2900 km) had shorter migration distances using the high point method (see below) compared to butterflies with pristine wings (3104 km; t=−2.13, p=0.04).
Butterflies were stored in glassine or paper envelopes and frozen or stored in desiccators until analysis. Wing tissue membrane was washed twice in 2:1 chloroform:methanol solution to remove surface oils and contaminants. Wing chitin subsamples (1.0±0.1 mg) for δ 13C were loaded into 8.0 mm×5.0 mm pressed tin capsules and combusted online using a Eurovector 3000 (Milan, Italy – www.eurovector.it) elemental analyzer. The resulting CO2 and N2 gases were separated by gas chromatography (GC) and introduced into a Nu Horizon (Nu Instruments, Wrexham, UK – www.nu-ins.com) triple-collector isotope-ratio mass-spectrometer via an open split and compared to a pure CO2 or N2 reference gas. Wing chitin subsamples (0.35±0.02 mg) for δ 2H isotopes were loaded into 4.0 mm×3.2 mm silver capsules, loaded into a zero-blank carousel and combusted using flash pyrolysis (1350 °C). Hydrogen gas was separated in a Eurovector Elemental Analyzer (Milan, Italy) and isotopic measurements made using an interfaced Elementar Isoprime™ (Isoprime, Manchester, UK) Continuous Flow Isotope Mass Spectrometer (CFIRMS). Non-exchangeable hydrogen δ 2H values for monarch wing chitin were obtained using the Comparative Equilibrium procedure [40], with results normalized to the VSMOW-SLAP scales. Laboratory keratin standards were equilibrated with the lab atmosphere for 72 h along with samples. The assigned values for hydrogen isotopes were CBS and KHS standards, currently having δ 2HVSMOW values of −197‰ and −54‰, respectively. Within-run reproducibility (n=5) for δ 2H values of keratin standards were better than±2‰. Laboratory standards for δ 13C were BWBII and PUGEL with assigned δ 13CVPDB values of −18.5‰ and −13.6‰ vs. the VPDB primary standard. The within-run precision of laboratory keratin control standards (n=5) were better than±0.15‰ for δ 13C. All analyses were conducted at the Stable Isotope Hydrology and Ecology Laboratory, Environment Canada, Saskatoon, Saskatchewan.
Migratory distance was defined as the distance between collection location at the overwintering colonies in Mexico and the isotopically assigned natal origin of each monarch using δ 13C and δ 2H measurements. Details of probabilistic assignment of natal origin are found in [32] but we briefly summarize the approach below. We used multivariate normal distribution assignment models to calculate the probability of natal origin to each pixel in a continuous landscape that spans the monarch breeding distribution [34, 36, 41]. For each butterfly, the model calculated the probability of natal origin to each pixel in our study area based on the correspondence between δ 2H and δ 13C values in wing tissue to the isoscape-predicted values of monarch δ 2H and δ 13C wing tissue of each geographically indexed cell in the landscape [42].
The probability density of individual i having location j as the natal origin is Y i ~ N(μ j,Σ) where Y i is a vector of observed δ 2H and δ 13C values, μ j is a vector of predicted δ 2H and δ 13C values derived from calibrated isoscapes [32], and Σ is the positive-definite variance-covariance matrix of δ 2H and δ 13C. Here, Σ was assumed to be constant across the entire isoscape and was estimated based on all values from known-location butterflies from data in [15]. We applied Bayes' rule to invert the conditional probabilities of natal origin based on isotopes as:
where fJ|Y,X,M is the spatially explicit posterior probability density function for location j as the true origin of individual with measured isotope value y, given the measured isotope values y ij for locations x j. The function fY|X represents the conditional distribution on Y j from above. The function fJ|M is the probability of occurrence for locations J, using a distributional prior of the breeding distribution, M, as described in [32].
From this probabilistic surface we calculated that latitude and longitude of natal origin in two ways. The high point method considered the latitude and longitude of the pixel with the highest probability to be the natal origin; we present the statistical results using the high point method in the main text of the manuscript. The centroid method calculated the mean latitude and mean longitude of the natal origin based on the odds that a given assigned origin was correct relative to the odds that it was incorrect as 2:1 and coded the upper 33% of the assignment surface for each butterfly as a binary surface [43]. The odds ratio constitutes the compromise between having sufficient geographic structure in the assignments while correctly assigning the natal origin of an individual [44], and is akin to choosing a type I error rate (e.g. α=0.05) in a traditional statistical test to determine significance. The centroid method accounts for uncertainty in the assignment of natal origin and reduces the potential effects of natal origins that are assigned near the edge of the breeding distribution based on a limited geographic extent of the isoscapes. The statistical results using the centroid method are presented in the supplementary material. Migration distance was calculated as a straight-line distance from the latitude and longitude of the assigned natal origins to the Sierra Chincua colony (19°40'20.6"N, 100°17'42.6"W; [45]).
Temperature-dependent development in monarchs may result in butterflies from northern areas developing at slower rates than southern areas [32]. If so, then controlling for temperature at the natal location in statistical analysis would allow us to account for environmental effects on wing morphology. Daily maximum temperature data for North America was downloaded [46, 47] and we calculated the mean maximum daily temperature between July 20 and August 9, which is the development period for overwintering butterflies when they are larvae (following Flockhart et al. [32]). For each butterfly, we extracted the mean maximum daily temperature value at both the high point and centroid location. To assess the effects of collinearity, we calculated the variance inflation factor for each variable in generalized linear models [48]. Variance inflation factors assess the boosted standard errors of parameters in generalized linear models which may obscure significant relationships between explanatory and response variables. The variance inflation factors of the variables in our models were generally lower than 3 (Additional file 1: Table S2) which is the recommended value when analysts should to take action to address collinearity [48] so we so ran analyses (see below) both with and without mean maximum daily temperature.
To examine how migration distance influenced wing morphology, we constructed global linear mixed effects models using the lme4 package [49] in program R for each response variable: forewing area, forewing length, roundness, and aspect ratio. These global models included sex and migration distance as fixed effects and year as a random effect. Sex was included because males had larger wings than females [26, 27]. We included year as a random effect because selection pressures likely change each year based on regional temperature-dependent development schedules that influence breeding distribution [36, 50–52] and autumn weather patterns that influence survival during migration, but these conditions were not expected to change in a predictable pattern over time. In a repeated analysis we also included mean maximum daily temperature as a fixed factor.
After constructing each global model, we removed the distance parameter for each response variable, and compared the reduced model to the global model using a likelihood ratio test [53]. This allowed us to evaluate whether the distance parameter significantly improved the model fit. We also ran the same series of global models as generalized linear models, substituting the Southern Oscillation Index (SOI) for year as a fixed effect in the model. SOI is a large-scale measure of climate that would influence all monarchs similarly in a given year and has a significant influence on the assigned natal origins of monarchs overwintering in Mexico [32]. Model outputs from the generalized linear models, as well as all models that include mean maximum daily temperature at the natal location, raw data, and R code are found in the Additional files 1, 2, and 3.
Monarchs in our study were estimated to have migrated between 774 km and 4430 km from natal locations to overwintering sites. Migratory distance was correlated with δ 2H values but this relationship was not linear (Additional file 1: Figure S1), suggesting that our two-isotope assignment approach was likely a better estimate of migration distance than using δ 2H values alone. Mean maximum daily temperature at natal origins ranged between 15.1–38.3 °C and, as expected, migratory distance and mean maximum daily temperature were correlated (high point: r=−0.78; centroid: r=−0.8; Additional file 1: Figure S1).
There were no significant differences between males and females in migration distance (t=−0.01, p=0.99). On average, male monarchs had longer forewings and greater forewing area than females, but males and females had similar wing roundness and aspect ratios (Additional file 1: Table S2). Using mixed-effects models that included year as a random effect (Additional file 1: Figure S2), both male and female monarchs that migrated longer distances had wings that were longer and had larger wing area (Table 1, Fig. 2). Although the parameter estimates were in the predicted direction, monarchs that migrated longer distances did not have more pointed wings or larger aspect ratios (Table 1). The direction of the parameter estimates and statistical inference between migratory distance and wing morphology was consistent when considering these relationships with generalized linear models that quantified annual variation using global oscillations indices (Additional file 1: Table S3). Global oscillation indices were not significant predictors of wing area (t=0.57, p=0.57), wing length (t=1.47, p=0.14) or aspect ratio (t=1.73, p=0.08) but were negatively related to wing roundness (β=−0.0008, t=3.17, p=0.001). Results using the high point method were consistent with the centroid method for estimating migration distance in both mixed-effects (Additional file 1: Table S4) and generalized linear models (Additional file 1: Table S5).
Variance inflation factors were generally less than 3 indicating that collinearity would have limited effects on drawing inference on the effects of migration distance on wing morphology (Additional file 1: Table S2). After controlling for mean maximum daily temperature, monarchs that migrated longer distances had wings that were longer and had larger wing area but there was no effect of migration distance on wing roundness and aspect ratio (Additional file 1: Tables S6). Using generalized linear models, there was no effect of mean maximum daily temperature on any wing attribute but the effect of migration distance on wing area and length remained significant (Additional file 1: Tables S7). There was no effect of migration distance on wing roundness or aspect ratio (Additional file 1: Tables S7). Similar to above, after controlling for mean maximum daily temperature the results using the high point method were consistent with the centroid method for estimating migration distance in both mixed-effects (Additional file 1: Table S8) and generalized linear models (Additional file 1: Table S9).
Consistent with previous studies [26, 28, 30, 31], our results showed that monarchs that migrated the farthest distance on average from natal grounds to wintering sites had the longest and largest wings. This observation suggests that migration acted as a selective episode on monarch butterfly wing morphology, specifically on wing length and area, and this relationship did not arise from temperature-dependent effects on wing morphology. Our study extended previous findings by proposing a mechanism by which natural selection may operate on wing morphology in migratory populations – longer and larger wings lead to higher survival of monarch butterflies during long-distance migration. In other words, monarchs that initiate fall migration from the most extreme limit of their range are more likely to successfully reach their wintering sites if they have longer wings and greater wing surface area, whereas shorter winged monarchs with small wing surface area have a greater chance of successful migration if they start from locations closer to the wintering sites. This result appeared to be consistent over the 40-year time series of monarchs collected in Mexico.
Efficient flight is important for small insects flying thousands of kilometers between breeding and non-breeding areas, and for which lipid accumulation during long distance movement is important for migration and overwinter survival [3, 54]. During migration, monarchs use both powered and gliding flight [55, 56]. Powered flight is more efficient with long wings and gliding/soaring flight is more efficient with reduced wing loading (ratio of mass:wing area; [13]). While we did not measure wing loading directly, we found a positive relationship between migration distance and forewing area, one component of wing loading. Moreover, prior work showed that wing area is strongly positively correlated to both body length (from head to abdomen) and total body mass of monarchs [26], suggesting that monarchs with larger wing area are also larger in body size overall. Thus, work here provides partial evidence that selection might operate on monarch wing length and area that influences wing loading during migration. This finding has also been reported for studies on migratory birds, which found a relationship between migration distance, wing morphology and body size at both the genus [7, 10] and species [8, 9, 11] levels. Skeletal morphology that promotes efficient travel has also been reported for salmon that migrated long distances [2], illustrating that selection for efficient travel during migration is important in non-flying migrants as well.
There was no evidence that wing roundness and aspect ratio correlated with migratory distance in our sample of monarchs, although the parameter estimates measured here were in the predicted direction of greater distance being associated with more angular (less round) wings. In a cross-population comparison, Davis and Altizer [26] found that non-migratory monarchs from Central America, Florida and the Caribbean have rounder (less elongated) forewings than migratory monarchs from eastern North America. Moreover, Dockx [27] showed that among monarchs captured in Cuba, migrants originating from northern latitudes had more angular wings than those identified as year-round residents. In more recent work, however, Li et al. [28] found no difference in wing shape among monarchs from migratory versus non-migratory populations, and Altizer et al. [30] also found no relationship between wing shape and estimated migration distance among eastern North American migrants. Thus, evidence for selection operating on wing shape in migratory monarchs appears to be mixed.
While evidence suggests that wing traits linked to migration in monarchs are genetically heritable [26], it is important to note that the fitness benefit of longer and larger wings could vary over different portions of the monarch's annual cycle or even after arriving at the overwintering colonies. Monarchs overwinter in dense colonies of millions of individuals with specific microclimatic conditions [57]. Clusters of butterflies retain heat, and monarchs in the middle of clusters are more sheltered from extreme temperatures and moisture on their wings that can freeze and kill them [57, 58]. Theoretically, larger monarchs might compete better for positions near the core of clusters; this advantage could be bolstered by earlier arrival times at the overwintering colonies. On the other hand, high conspecific density in colonies [up to 50 million butterflies per hectare [57] may result in large individuals having a disadvantage for obtaining an optimal roosting location, which requires precise and controlled flight in confined spaces. Smaller males are also more likely to mate at the overwintering sites than larger males [59], although this appears to be caused more by small males initiating mating attempts as opposed to a selective advantage conferred by small wings, as captive experiments showed that males with larger wings mated more often than small males [60]. Additionally, monarchs breed over successive generations [36, 51] and summer migratory distances must be significantly shorter than the fall migration distances and hence selection may not be as strong during the breeding season. Wing measurements of butterflies collected during the breeding season coupled with migratory distance estimates derived from stable isotopes would inform whether selection operates during the breeding season.
Individuals with larger wings might experience lower wing loading and greater flight efficiency during long-distance migration [12, 61]. On the other hand, Borland et al. [62] found no difference in the wing length of butterflies captured during migration through the USA in Minnesota versus Texas, which suggests that if smaller individuals were dying during migration, this happened after monarchs entered Mexico [54]. If so, then we would also predict that monarchs that remigrate from Mexico to Texas to breed would also have longer wings than those migrating through Texas the previous autumn. Given the prediction that longer wings should increase survival of monarchs during migration, we would also predict that larger butterflies could amass great fat reserves to fuel the long overwintering period. Monarchs in particular accumulate fat reserves during that last portion of migration between Texas and overwintering colonies in Mexico [63]. From this perspective, there could be a complex interaction among wing loading, lipid reserves and successful migration.
Although results here indicate that long-distance migration selects for larger wings, we are not suggesting that this will necessarily result in the evolution of longer-winged sub-populations within the eastern migratory population. Since monarchs of all migratory distances overwinter together each winter and breed in the spring, any effect of wing size selection will be limited by population mixing during migration itself and at the overwintering sites [22, 31, 64–66]. Moreover, because of the multiple-generation aspect of monarch migration, individuals do not usually return and breed in their natal region [35], making it unlikely that the same selection pressures will act on the genetic line of longer-distance migrants in the following migratory season. Therefore, although the migratory population in eastern North America is morphologically different from non-migratory ones in other parts of the world, it is unlikely that eastern monarchs will experience directional selection on wing morphology. The recurring selective disappearance of individuals with shorter wings during autumn migration arises from a dilution of the selection pressures acting on wing morphology across multiple breeding generations. If this dilution effect operates in other multivoltine insects, then migratory behaviour may be less likely to evolve in insects compared to vertebrate taxa.
Understanding the determinants of successful migration during the annual cycle could improve understanding of how populations will respond to future conditions. For example, over the coming century, the distribution of both monarchs and their host plants are predicted to move northward due to climate change [67, 68]. If this bears out, these distributional changes will require individuals to successfully complete longer autumn migrations to traditional overwintering areas in Mexico. The prospect that individuals with longer wings have an advantage during long-distance migration suggests that at least some monarchs should be able to conduct this longer migration to overwintering locations in Mexico in the future. Although monarchs may colonize and successfully breed in more northern areas in coming decades, biologists should not assume that that will automatically enhance or stabilize population persistence if these same morphologies are more strongly selected against at different portions of the annual cycle. Future work might well consider how competing selective forces of flight maneuverability, thermoregulation, and migration distance interact to influence wing morphology and the consequences for population distribution of the monarch butterfly.
Additional results. (DOCX 6879 kb)
The data set used in the analysis. (XLSX 801 bytes)
The R code to run the analysis. (DOCX 14 kb)
Geoff Koehler assisted with stable isotope measurements at the NHRC laboratory of Environment and Climate Change Canada.
All isotopic measurements were performed at Environment Canada in Saskatoon, Canada, and financed in part from operating grants from Environment Canada to KAH and LIW. DTTF was supported by a Liber Ero postdoctoral fellowship. LPB's butterfly samples were collected during research trips supported by the U.S. National Science Foundation, the Wildlife Conservation Society, the World Wildlife Fund, the October Hill Foundation, and Scion Natural Science Association. Samples from SA were supported by a grant from the National Science Foundation (DEB-064383). Funding agencies had no input on the design of the study and collection, analysis, and interpretation of the data and writing of the manuscript.
Data and the R code to run the models are available in the Additional Files.
LPB, SA, KAH, LIW, DTTF and DRN collected the samples. BF and RD measured the samples and prepared the samples for stable isotope analysis. KAH and LIW conducted the stable isotope analysis. BF, DTTF, RD and DRN conducted the statistical analysis and drafted the paper. All authors provided extensive revisions on the manuscript. All authors approve the submitted version of the manuscript.
The authors declare that they have no competing interests.
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No animal care approval was required for this research.
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D. T. Tyler Flockhart, Email: ac.hpleugou@ahkcolfd.
Blair Fitz-gerald, Email: ac.hpleugou.liam@10gztifb.
Lincoln P. Brower, Email: ude.cbs@reworb.
Rachael Derbyshire, Email: ac.hpleugou@hsybredr.
Sonia Altizer, Email: ude.agu@rezitlas.
Keith A. Hobson, Email: ac.owu@6nosbohk.
Leonard I. Wassenaar, Email: email@example.com.
D. Ryan Norris, Email: ac.hpleugou@sirronr.
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