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Majority consensus maximum likelihood tree (250 bootstrap replicates) based on comparison of all analyzed protein markers, showing the relationships between the analyzed spounaviruses.Branch labels indicate their percent bootstrap support. Leaves are colored by proposed in-subfamily clustering: blue ? Bastille group, green ? Twort group, red ? Bacillus phage SPO1. Abbreviations include name of host taxon (Ba ? Bacillus, Bx ? Brochothrix, En ? Enterococcus, Lb ? Lactobacillus, Li ? Listeria, St ? Staphylococcus) and the bacteriophage name. All analyzed sequences are listed in Table S1.
Webpy Howto
For the moment I'm going to try using Webpy. Why?
Well I like the fact that Aaron could get an app up quickly without having to make lots of adjustments to existing code. I'm not so sure this is the case with django. Which bits can you use? What bits are being left out?
I suppose the biggest problem I have with django is the size of the codebase you have to upload. More code, more moving parts and the greater chance for something to go wrong. Plus the fact you have to use a hack to even start (django_bootstrap.py). I don't think this is the real problem though. The real problem starts when you have to set up your local application and try to get it to run on appengine. There is a couple of little tricks & problems I've found. The approach is far from seamless and prone to error that will compound adding large frameworks. Not a show stopper but annoying enough to slow you down. The google developers are really pulling the stops out for django support so don't assume not choosing django now will stop me using it in the future.
Problems
The following solutions below should give you enough information to get webpy up and running. If you have an questions thow a question to this google app-engine thread, twitter.com/bootload or email me. Below is a list of the problems you will encounter.
1) Cannot run webpy locally:
I've encountered a few problems. The first is "how do you run webpy locally?" The problem is you may have webpy installed, the webpy code (web tree) in your current directory. But how does your local code find it? [1] Hidden in the documentation is a hint to make a symbolic link to the code you wish to use. Here is how:
* We wish to use Webpy and "import web"
* make sure you have a copy of "bzr" (debian/ubuntu): "apt-get install bzr"
* download the latest (0.3) webpy install version (0.3): "bzr get webpy.org/bzr/webpy.dev/;
* install the latest webpy, cd into webpy.dev & install: "python setup.py install"
* in the directory you have installed your app-engine client, create a symbolic link to a newly created directory called "web" to your webpy installation.
* ln -s /usr/lib/python2.5/site-packages/web web
The result is something like this with the app-engine code, your code (helloworld & hellowebpy).
drwxr-xr-x 12 fb fb 4096 2008-04-18 20:09 .
drwxr-xr-x 18 fb fb 4096 2008-04-14 21:23 ..
-r-xr-xr-x 1 fb fb 1595 2008-04-03 11:05 appcfg.py
-r--r--r-- 1 fb fb 154 2008-04-03 11:05 BUGS
drwxr-xr-x 4 fb fb 4096 2008-04-16 22:16 demos
-r-xr-xr-x 1 fb fb 1713 2008-04-16 16:59 dev_appserver.py
drwxr-xr-x 5 fb fb 4096 2008-04-14 22:05 google
drwxr-xr-x 8 fb fb 4096 2008-04-03 11:05 google_appengine
drwxr-xr-x 6 fb fb 4096 2008-04-18 11:47 hellowebpy
drwxr-xr-x 6 fb fb 4096 2008-04-16 15:57 helloworld
drwxr-xr-x 5 fb fb 4096 2008-04-03 11:05 lib
-r--r--r-- 1 fb fb 4348 2008-04-03 11:05 LICENSE
drwxr-xr-x 2 fb fb 4096 2008-04-09 13:55 new_project_template
-r--r--r-- 1 fb fb 3476 2008-04-03 11:05 README
drwxr-xr-x 2 fb fb 4096 2008-04-09 13:55 templates
drwxr-xr-x 2 fb fb 4096 2008-04-09 13:55 tools
-r--r--r-- 1 fb fb 57 2008-04-03 11:05 VERSION
lrwxrwxrwx 1 fb fb 36 2008-04-16 23:11 web -> /usr/lib/python2.5/site-packages/web
This now allows you to run your webpy code on the development machine.
2) Run webpy remotely :
In your source code directory (in my case, hellowebpy) take a copy of the webpy.dev/web directory and paste it into hellowebpy/. The result is something like this:
drwxr-xr-x 6 fb fb 4096 2008-04-18 20:24 .
drwxr-xr-x 12 fb fb 4096 2008-04-18 20:09 ..
-rw-r--r-- 1 fb fb 223 2008-04-18 11:05 app.yaml
-rwxr--r-- 1 fb fb 844 2008-04-13 23:23 favicon.ico
-rw-r--r-- 1 fb fb 448 2008-04-18 11:47 hello.py
drwxr-xr-x 2 fb fb 4096 2008-04-18 11:07 images
-rw-r--r-- 1 fb fb 471 2008-04-18 11:47 index.yaml
drwxr-xr-x 2 fb fb 4096 2008-04-18 11:07 styles
drwxr-xr-x 2 fb fb 4096 2008-04-18 10:57 templates
drwxr-xr-x 4 fb fb 4096 2008-04-18 11:08 web
* cd up to the root directory with appcfg.py
* upload your code + the webpy "web" code using appcfg.py: "./appcfg --noisy -e foo@bar.com update hellowebpy/" (remember to use your email address)
3) Problems with errors and c-based code :
If you have installed code like Cheetah you will get an error saying you cannot run c-based code. The reason is the dev_appserver.py code assumes the Cheetah code you have is trying to make a call to some cPython code, strop (a C based, string library optomised for speed). [3] The solution is to un-install Cheetah from your system. The google app-engine code base has a modified version of Cheetah removing all C code based routines. Be warned.
4) Problems uploading new code? :
I've had problems updating code sometimes. I suspect caching is the problem. So go to your root directory in your account and search for ".app*". You should see the following files, .appcfg_cookies and .appcfg_nag. Delete them both. Create a new directory, delete the index.yaml and copy your old code into the new directory. Then re-run the upload sequence ... "./appcfg --noisy -e foo@bar.com update new_directory_with_your_code/"
Following these steps should allow you to run webpy. Remember the example Aaron shows is for version 0.3. It was mentioned in the google-appengine webpy post others managed to get 0.23 to work. Try what they suggest. The example code works as shown for 0.3.
Have fun.
Reference
[1] Google App Engine, Pure Python, "... You can include other pure Python libraries with your application by putting the code in your application directory. If you make a symbolic link to a module's directory in your application directory, appcfg.py will follow the link and include the module in your app ..."
code.google.com/appengine/docs/python/purepython.html
[2] Python docs, "common string manipulations, optimized for speed called by "import string" /usr/local/lib/python2.4/lib-dynload/strop.so"
<<< start
Made with a Anne Marie Horner lawn (the stripe) and a lawn by Cotton + Steel purchased from www.fabric.com/buy/0395399/cotton-steel-tokyo-train-ride-....
Faux Suede on the inside collar - easier and itch free against the neck. The colors were a great match...can barely tell the difference in this photo, but the suede is a tad, just a scooch, lighter.
Covered buttons are the Dritz kits, machine button holes.
My friend Abi gifted me this yardage; I sewed it up on my grandmother's 1950 Singer 15-91.
I've blogged a little about construction; I am proud of this coat. It is underlined in flannel and features five hidden pockets, a padded hem, bust and back waist darts, and a bias-blocked sleeve.
My grandmother personified both a mature and rugged personal style, as well as a pirate-mouth! :) I miss her very much.
一小時 RWD 就上手(SUSY responsive grid for compass.)
(怎麼課程有三小時?因為要讓你練習啊!)
如果你
使用 bootstrap/foundation grid 之後,才發現它的 responsive grid 設定很腦殘。
想要控制每個中斷點(breakpoint)的欄位設計,而不只是一起縮小寬度或取消欄位而已。
想學習一套 mobile first 版面設計的最佳實踐方式。
那你就該來玩玩SUSY!
當全天下了無新意的使用 bootstrap/foundation 之後,聰明人就該跳出來自己設計網站了。其實 CSS3 的 media query 只是一個規格書,該如何好好利用做出任意又靈活的 Responsive Web Design,從 SUSY 入門是一個非常棒的選擇,從初學到客製化排版系統,一小時就能上手。
Here with my Vancouver homeboys from outsourcingthingsdone.com checking out the new crop of startups.
Perfect collaboration between bootstrap, xDetailView, GroupGridView in real accouting application.
it is just too wonderful....
Identification of variant XMRV in prostate cancer patients using phylogenetic analysis.
A. envelope (env), B. polymerase (pol), and C. gag. Stability of the tree topology was tested using 1000 bootstrap replicates in both neighbor joining (NJ) and maximum likelihood (ML) methods. Bootstrap values >60 are shown at major nodes (NJ/ML). New sequences from the current study are boxed. Accession numbers for prototypical MLV sequences available at GenBank are XMRV VP35 = DQ241301, XMRV VP62 = DQ399707, XMRV VP42 = DQ241302, XMRV WPI-1106 = GQ497344, XMRV WPI-1178 = GC497343, XMRV PCA1–PCA17 = GU812341–GU812357, MLV DG-75 = AF221065, MLV MTCR = NC_001702, MLV AKV = J01998, MLV BM5eco = AY252102.1, Moloney MLV = J02255, Moloney neuropthogenic MLV variant ts1-92b = AF462057, Rauscher MLV = NC_001819, Friend MLV = X02794, mERV Chr 7 = AC167978, mERV Chr 7 = AC127565, mERV Chr 8 = AC127575, mERV Chr 12 = AC153658, mERV Chr 9 = AC121813, mERV Chr 4 = AL627077, mERV Chr 1 = AC083892), XMLV A2780 = FR670594, XMLV BHY = FR670595, XMLV Daudi = FR670596, XMLV EKVX = FR670597, XMLV IMR-5 = FR670598, XMLV MUTZ-1 = FR670599, XMLV S-117 = FR670600, XMLV TYK-nu = FR670601. Sequences denoted RAW are from the polytropic MLV isolated in HeLa cells used to develop the in-house WB test. Sequences coded as XMRV VP and PCA and WPI are from prostate cancer and CFS patients, respectively. Additional prostate cancer patient VP gag and pol sequences were kindly provided by Drs. Robert Silverman and Joe Derisi. Viral tropism, as determined by analysis of env sequences, is indicated by blue (xenotropic), purple (polytropic), and yellow (ecotropic) spheres.
一小時 RWD 就上手(SUSY responsive grid for compass.)
(怎麼課程有三小時?因為要讓你練習啊!)
如果你
使用 bootstrap/foundation grid 之後,才發現它的 responsive grid 設定很腦殘。
想要控制每個中斷點(breakpoint)的欄位設計,而不只是一起縮小寬度或取消欄位而已。
想學習一套 mobile first 版面設計的最佳實踐方式。
那你就該來玩玩SUSY!
當全天下了無新意的使用 bootstrap/foundation 之後,聰明人就該跳出來自己設計網站了。其實 CSS3 的 media query 只是一個規格書,該如何好好利用做出任意又靈活的 Responsive Web Design,從 SUSY 入門是一個非常棒的選擇,從初學到客製化排版系統,一小時就能上手。
Find the top Bootstrap themes that are responsive for mobile and
tablets. Stunning website graphic designs using the Bootstrap layout..For more information to log in my website codeshop.co
Maximum likelihood phylogeny of ?9 desaturases in only social Hymenoptera.Bootstrap values greater than 50 are shown on the tree. Branches with a bootstrap value under 50 are collapsed to show unresolved parts of the tree and underline supported clades. Clades shown with vertical red bars correspond to well-supported clades chosen to perform selection analysis. Non-ant social insect desaturase sequences are shown in pink (Bter: Bombus terrestris; Bimp: Bombus impatiens; Amel: Apis mellifera; Aflo: Apis floridanus). Ant desaturase sequences are shown in red (Hsal: Harpegnathos saltator; Lhum: Linepithema humile; Cflo: Camponotus floridanus; Fexs: Formica exsecta; Pbar: Pogonomyrmex barbatus; Acep: Atta cephalotes; Aech: Acromymex echinatior; Sinv: Solenopsis invicta).
pictionid57358521 - catalog14039451 - titleatlas 72d details missile 72d lox bootstrap flexline date 03031966 - filename14039451.jpgImages from the Convair/General Dynamics Astronautics Atlas Negative Collection. The processing, cataloging and digitization of these images has been made possible by a generous National Historical Publications and Records grant from the National Archives and Records Administration---Please Tag these images so that the information can be permanently stored with the digital file.---Repository: San Diego Air and Space Museum
一小時 RWD 就上手(SUSY responsive grid for compass.)
(怎麼課程有三小時?因為要讓你練習啊!)
如果你
使用 bootstrap/foundation grid 之後,才發現它的 responsive grid 設定很腦殘。
想要控制每個中斷點(breakpoint)的欄位設計,而不只是一起縮小寬度或取消欄位而已。
想學習一套 mobile first 版面設計的最佳實踐方式。
那你就該來玩玩SUSY!
當全天下了無新意的使用 bootstrap/foundation 之後,聰明人就該跳出來自己設計網站了。其實 CSS3 的 media query 只是一個規格書,該如何好好利用做出任意又靈活的 Responsive Web Design,從 SUSY 入門是一個非常棒的選擇,從初學到客製化排版系統,一小時就能上手。
Made with a Anne Marie Horner lawn (the stripe) and a lawn by Cotton + Steel purchased from www.fabric.com/buy/0395399/cotton-steel-tokyo-train-ride-....
一小時 RWD 就上手(SUSY responsive grid for compass.)
(怎麼課程有三小時?因為要讓你練習啊!)
如果你
使用 bootstrap/foundation grid 之後,才發現它的 responsive grid 設定很腦殘。
想要控制每個中斷點(breakpoint)的欄位設計,而不只是一起縮小寬度或取消欄位而已。
想學習一套 mobile first 版面設計的最佳實踐方式。
那你就該來玩玩SUSY!
當全天下了無新意的使用 bootstrap/foundation 之後,聰明人就該跳出來自己設計網站了。其實 CSS3 的 media query 只是一個規格書,該如何好好利用做出任意又靈活的 Responsive Web Design,從 SUSY 入門是一個非常棒的選擇,從初學到客製化排版系統,一小時就能上手。
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matching people to places)
Here with my Vancouver homeboys from outsourcingthingsdone.com checking out the new crop of startups.
Here with my Vancouver homeboys from outsourcingthingsdone.com checking out the new crop of startups.
一小時 RWD 就上手(SUSY responsive grid for compass.)
(怎麼課程有三小時?因為要讓你練習啊!)
如果你
使用 bootstrap/foundation grid 之後,才發現它的 responsive grid 設定很腦殘。
想要控制每個中斷點(breakpoint)的欄位設計,而不只是一起縮小寬度或取消欄位而已。
想學習一套 mobile first 版面設計的最佳實踐方式。
那你就該來玩玩SUSY!
當全天下了無新意的使用 bootstrap/foundation 之後,聰明人就該跳出來自己設計網站了。其實 CSS3 的 media query 只是一個規格書,該如何好好利用做出任意又靈活的 Responsive Web Design,從 SUSY 入門是一個非常棒的選擇,從初學到客製化排版系統,一小時就能上手。
Here with my Vancouver homeboys from outsourcingthingsdone.com checking out the new crop of startups.
Third part of three (Figs. 3, 4 and 5): Phylogenetic tree of nifH protein sequences.50% majority rule consensus tree of 13,500 PhyloBayes [52] post burn-in trees, unrooted. Black values at internodes ?=? Bayesian Posterior Probability (if >0.5). Pink values ?=? MEGA5 [56] Maximum Parsimony (MP) bootstrap support (if >50). Green values ?=? GARLI [54] Maximum Likelihood bootstrap support (if >50). Terminal triangles represent monophyletic clades with MOTUs solely of one tree species origin, collapsed but keeping the internal distance (substitutions per site, see scale bar), in light pink ?=? 50?79 MP bootstrap support, dark pink ?=? 80?100 MP bootstrap support. Green color indicates MOTUs solely from Fagus origin, red color Picea origin and dark blue color mixed origin (with bars showing ratio of [green] vs. [red]). Terminal labels with sequences from this study: MOTU ID (SMOTU ?=? singleton MOTU), total number of sequences, FASY ?=? from Fagus, PIAB ?=? from Picea, followed by number of sequences in the same order, then forest management type(s) (AC.Conif ?=? managed spruce forests, AC.Decid ?=? managed beech forests, Extensiv ?=? extensively managed beech forests) and number of sequences in same order. Terminal labels with sequences from other sources: near BLAST hit, summary of ecological data of sequences in that MOTU. The width of visible terminal branches represents the number of sequences (size correct up to 10 sequences). To the right, amino acid sequence logos and Kyte-Doolittle hydophobicity alignments for labeled nodes on the tree. The small tree shape (based on screenshot from Archaeopteryx v.0.972 [66]) shows the position within the complete phylogenetic tree.
Made my simple quotes app responsive and flat html, performant, simple, mobile and web... best of all worlds.
Here with my Vancouver homeboys from outsourcingthingsdone.com checking out the new crop of startups.
Clear is a vue or Vuejs based admin template built with bootstrap, also comes with laravel version. It utilizes vuex, vue-router and comes with laravel spark skin.
Find the top Bootstrap themes that are responsive for mobile and
tablets. Stunning website graphic designs using the Bootstrap layout..For more information to log in my website codeshop.co
Scott B Reynolds gave a presentation on designing e-commerce websites. The discussion included Adobe Dreamweaver, Business Catalyst, and Twitter's Bootstrap v2. Hartford Adobe
一小時 RWD 就上手(SUSY responsive grid for compass.)
(怎麼課程有三小時?因為要讓你練習啊!)
如果你
使用 bootstrap/foundation grid 之後,才發現它的 responsive grid 設定很腦殘。
想要控制每個中斷點(breakpoint)的欄位設計,而不只是一起縮小寬度或取消欄位而已。
想學習一套 mobile first 版面設計的最佳實踐方式。
那你就該來玩玩SUSY!
當全天下了無新意的使用 bootstrap/foundation 之後,聰明人就該跳出來自己設計網站了。其實 CSS3 的 media query 只是一個規格書,該如何好好利用做出任意又靈活的 Responsive Web Design,從 SUSY 入門是一個非常棒的選擇,從初學到客製化排版系統,一小時就能上手。
Here with my Vancouver homeboys from outsourcingthingsdone.com checking out the new crop of startups.
一小時 RWD 就上手(SUSY responsive grid for compass.)
(怎麼課程有三小時?因為要讓你練習啊!)
如果你
使用 bootstrap/foundation grid 之後,才發現它的 responsive grid 設定很腦殘。
想要控制每個中斷點(breakpoint)的欄位設計,而不只是一起縮小寬度或取消欄位而已。
想學習一套 mobile first 版面設計的最佳實踐方式。
那你就該來玩玩SUSY!
當全天下了無新意的使用 bootstrap/foundation 之後,聰明人就該跳出來自己設計網站了。其實 CSS3 的 media query 只是一個規格書,該如何好好利用做出任意又靈活的 Responsive Web Design,從 SUSY 入門是一個非常棒的選擇,從初學到客製化排版系統,一小時就能上手。
My friend Abi gifted me this yardage; I sewed it up on my grandmother's 1950 Singer 15-91.
I've blogged a little about construction; I am proud of this coat. It is underlined in flannel and features five hidden pockets, a padded hem, bust and back waist darts, and a bias-blocked sleeve.
My grandmother personified both a mature and rugged personal style, as well as a pirate-mouth! :) I miss her very much.
My friend Abi gifted me this yardage; I sewed it up on my grandmother's 1950 Singer 15-91.
I've blogged a little about construction; I am proud of this coat. It is underlined in flannel and features five hidden pockets, a padded hem, bust and back waist darts, and a bias-blocked sleeve.
My grandmother personified both a mature and rugged personal style, as well as a pirate-mouth! :) I miss her very much.
Phylogenetic relationships among haplotypes and lineage subdivergence detected in Primula obconica.(a) The phylogenetic topography based on plastid DNA dataset. Bootstrap values of Maximum Parsimony analysis and posterior probabilities of Bayesian inference are given above and below branches, respectively. (b) Maximum Parsimony networks of chlorotypes identified by TCS. Each solid line between circles represents one mutational step between two chlorotypes based on most parsimonious algorithm. The small open circles indicate the missing chlorotypes (not sampled or extinct). The solid line in the middle position of the network represents the two main lineages identified in the phylogenetic analysis, while the dashed line indicates the subdivision in each main lineage. The arrow indicated the connection between Primua obconica and Primula barbicalyx. Yellow and green circles in (a) and (b) correspond to lineage A and lineage B, respectively, as shown in Figure 1. (c) The strict consensus of the Maximum Parsimony trees of ribotypes. The two main lineages are circled by a solid line, while a dashed line in each circle represents the subdivision in each main lineage. The terminal of each branch represents haplotype recovered from plastid DNA and ITS datasets (See Table 1 and 3).
Bats are mammals of the order Chiroptera (/kaɪˈrɒptərə/; from the Greek χείρ - cheir, "hand" and πτερόν - pteron, "wing") whose forelimbs form webbed wings, making them the only mammals naturally capable of true and sustained flight. By contrast, other mammals said to fly, such as flying squirrels, gliding possums, and colugos, can only glide for short distances. Bats do not flap their entire forelimbs, as birds do, but instead flap their spread-out digits, which are very long and covered with a thin membrane or patagium.
Bats are the second largest order of mammals (after the rodents), representing about 20% of all classified mammal species worldwide, with about 1,240 bat species divided into two suborders: the less specialized and largely fruit-eating megabats, or flying foxes, and the highly specialized and echolocating microbats. About 70% of bat species are insectivores. Most of the rest are frugivores, or fruit eaters. A few species, such as the fish-eating bat, feed from animals other than insects, with the vampire bats being hematophagous, or feeding on blood.
Bats are present throughout most of the world, with the exception of extremely cold regions. They perform vital ecological roles of pollinating flowers and dispersing fruit seeds; many tropical plant species depend entirely on bats for the distribution of their seeds. Bats are economically important, as they consume insect pests, reducing the need for pesticides. The smallest bat is the Kitti's hog-nosed bat, measuring 29–34 mm in length, 15 cm across the wings and 2–2.6 g in mass. It is also arguably the smallest extant species of mammal, with the Etruscan shrew being the other contender. The largest species of bat are a few species of Pteropus (fruit bats or flying foxes) and the giant golden-crowned flying fox with a weight up to 1.6 kg and wingspan up to 1.7 m.
CLASSIFICATION AND EVOLUTION
Bats are mammals. In many languages, the word for "bat" is cognate with the word for "mouse": for example, chauve-souris ("bald-mouse") in French, murciélago ("blind mouse") in Spanish, saguzahar ("old mouse") in Basque, летучая мышь ("flying mouse") in Russian, slijepi miš ("blind mouse") in Bosnian, nahkhiir ("leather mouse") in Estonian, vlermuis (winged mouse) in Afrikaans, from the Dutch word vleermuis (from Middle Dutch "winged mouse"). An older English name for bats is flittermouse, which matches their name in other Germanic languages (for example German Fledermaus and Swedish fladdermus). Bats were formerly thought to have been most closely related to the flying lemurs, treeshrews, and primates, but recent molecular cladistics research indicates that they actually belong to Laurasiatheria, a diverse group also containing Carnivora and Artiodactyla.
The two traditionally recognized suborders of bats are:
- Megachiroptera (megabats)
- Microchiroptera (microbats/echolocating bats)
Not all megabats are larger than microbats. The major distinctions between the two suborders are:
- Microbats use echolocation; with the exception of the Rousettus genus, megabats do not.
- Microbats lack the claw at the second finger of the forelimb.
- The ears of microbats do not close to form a ring; the edges are separated from each other at the base of the ear.
- Microbats lack underfur; they are either naked or have guard hairs.
Megabats eat fruit, nectar, or pollen. Most microbats eat insects; others may feed on fruit, nectar, pollen, fish, frogs, small mammals, or the blood of animals. Megabats have well-developed visual cortices and show good visual acuity, while microbats rely on echolocation for navigation and finding prey.
The phylogenetic relationships of the different groups of bats have been the subject of much debate. The traditional subdivision between Megachiroptera and Microchiroptera reflects the view that these groups of bats have evolved independently of each other for a long time, from a common ancestor already capable of flight. This hypothesis recognized differences between microbats and megabats and acknowledged that flight has only evolved once in mammals. Most molecular biological evidence supports the view that bats form a single or monophyletic group.
Researchers have proposed alternative views of chiropteran phylogeny and classification, but more research is needed.
In the 1980s, a hypothesis based on morphological evidence was offered that stated the Megachiroptera evolved flight separately from the Microchiroptera. The so-called flying primates theory proposes that, when adaptations to flight are removed, the Megachiroptera are allied to primates by anatomical features not shared with Microchiroptera. One example is that the brains of megabats show a number of advanced characteristics that link them to primates. Although recent genetic studies strongly support the monophyly of bats, debate continues as to the meaning of available genetic and morphological evidence.
Genetic evidence indicates that megabats originated during the early Eocene and should be placed within the four major lines of microbats.
Consequently, two new suborders based on molecular data have been proposed. The new suborder of Yinpterochiroptera includes the Pteropodidae, or megabat family, as well as the Rhinolophidae, Hipposideridae, Craseonycteridae, Megadermatidae, and Rhinopomatidae families The other new suborder, Yangochiroptera, includes all of the remaining families of bats (all of which use laryngeal echolocation). These two new suborders are strongly supported by statistical tests. Teeling (2005) found 100% bootstrap support in all maximum likelihood analyses for the division of Chiroptera into these two modified suborders. This conclusion is further supported by a 15-base-pair deletion in BRCA1 and a seven-base-pair deletion in PLCB4 present in all Yangochiroptera and absent in all Yinpterochiroptera. Perhaps most convincingly, a phylogenomic study by Tsagkogeorga et al (2013) showed that the two new proposed suborders were supported by analyses of thousands of genes.
The chiropteran phylogeny based on molecular evidence is controversial because microbat paraphyly implies that one of two seemingly unlikely hypotheses occurred. The first suggests that laryngeal echolocation evolved twice in Chiroptera, once in Yangochiroptera and once in the rhinolophoids. The second proposes that laryngeal echolocation had a single origin in Chiroptera, was subsequently lost in the family Pteropodidae (all megabats), and later evolved as a system of tongue-clicking in the genus Rousettus.
Analyses of the sequence of the "vocalization" gene, FoxP2, were inconclusive as to whether laryngeal echolocation was secondarily lost in the pteropodids or independently gained in the echolocating lineages. However, analyses of the "hearing" gene, Prestin seemed to favor the independent gain in echolocating species rather than a secondary loss in the pteropodids.
In addition to Yinpterochiroptera and Yangochiroptera, the names Pteropodiformes and Vespertilioniformes have also been proposed for these suborders. Under this new proposed nomenclature, the suborder Pteropodiformes includes all extant bat families more closely related to the genus Pteropus than the genus Vespertilio, while the suborder Vespertilioniformes includes all extant bat families more closely related to the genus Vespertilio than to the genus Pteropus.
Little fossil evidence is available to help map the evolution of bats, since their small, delicate skeletons do not fossilize very well. However, a Late Cretaceous tooth from South America resembles that of an early microchiropteran bat. Most of the oldest known, definitely identified bat fossils were already very similar to modern microbats. These fossils, Icaronycteris, Archaeonycteris, Palaeochiropteryx and Hassianycteris, are from the early Eocene period, 52.5 million years ago. Archaeopteropus, formerly classified as the earliest known megachiropteran, is now classified as a microchiropteran.
Bats were formerly grouped in the superorder Archonta, along with the treeshrews (Scandentia), colugos (Dermoptera), and the primates, because of the apparent similarities between Megachiroptera and such mammals. Genetic studies have now placed bats in the superorder Laurasiatheria, along with carnivorans, pangolins, odd-toed ungulates, even-toed ungulates, and cetaceans. A recent study by Zhang et al. places Chiroptera as a sister taxon to the clade Perissodactyla (which includes horses and other odd-toed ungulates). However, the first phylogenomic analysis of bats shows that they are not sisters to Perissodactyla, instead they are sisters to a larger group that includes ungulates and carnivores.
Megabats primarily eat fruit or nectar. In New Guinea, they are likely to have evolved for some time in the absence of microbats, which has resulted in some smaller megabats of the genus Nyctimene becoming (partly) insectivorous to fill the vacant microbat ecological niche. Furthermore, some evidence indicates that the fruit bat genus Pteralopex from the Solomon Islands, and its close relative Mirimiri from Fiji, have evolved to fill some niches that were open because there are no nonvolant or nonflying mammals on those islands.
FOSSIL BATS
Fossilized remains of bats are few, as they are terrestrial and light-boned. Only an estimated 12% of the bat fossil record is complete at the genus level. Fossil remains of an Eocene bat, Icaronycteris, were found in 1960. Another Eocene bat, Onychonycteris finneyi, was found in the 52-million-year-old Green River Formation in Wyoming, United States, in 2003. This intermediate fossil has helped to resolve a long-standing disagreement regarding whether flight or echolocation developed first in bats. The shape of the rib cage, faceted infraspious fossa of the scapula, manus morphology, robust clavicle, and keeled sternum all indicated Onychonycteris was capable of powered flight. However, the well-preserved skeleton showed that the small cochlea of the inner ear did not have the morphology necessary to echolocate. O. finneyi lacked an enlarged orbical apophysis on the malleus, and a stylohyal element with an expanded paddle-like cranial tip - both of which are characteristics linked to echolocation in other prehistoric and extant bat species. Because of these absences, and the presence of characteristics necessary for flight, Onychonycteris provides strong support for the “flight first” hypothesis in the evolution of flight and echolocation in bats.
The appearance and flight movement of bats 52.5 million years ago were different from those of bats today. Onychonycteris had claws on all five of its fingers, whereas modern bats have at most two claws appearing on two digits of each hand. It also had longer hind legs and shorter forearms, similar to climbing mammals that hang under branches such as sloths and gibbons. This palm-sized bat had short, broad wings, suggesting it could not fly as fast or as far as later bat species. Instead of flapping its wings continuously while flying, Onychonycteris likely alternated between flaps and glides while in the air. Such physical characteristics suggest that this bat did not fly as much as modern bats do, rather flying from tree to tree and spending most of its waking day climbing or hanging on the branches of trees. The distinctive features noted on the Onychonycteris fossil also support the claim that mammalian flight most likely evolved in arboreal gliders, rather than terrestrial runners. This model of flight development, commonly known as the "trees-down" theory, implies that bats attained powered flight by taking advantage of height and gravity, rather than relying on running speeds fast enough for a ground-level take off.
The mid-Eocene genus Necromantis is one of the earliest examples of bats specialised to hunt vertebrate prey, as well as one of the largest bats of its epoch.
HABITATS
Flight has enabled bats to become one of the most widely distributed groups of mammals. Apart from the Arctic, the Antarctic and a few isolated oceanic islands, bats exist all over the world. Bats are found in almost every habitat available on Earth. Different species select different habitats during different seasons, ranging from seasides to mountains and even deserts, but bat habitats have two basic requirements: roosts, where they spend the day or hibernate, and places for foraging. Most temperate species additionally need a relatively warm hibernation shelter. Bat roosts can be found in hollows, crevices, foliage, and even human-made structures, and include "tents" the bats construct by biting leaves.
The United States is home to an estimated 45 to 48 species of bats. The three most common species are Myotis lucifugus (little brown bat), Eptesicus fuscus (big brown bat), and Tadarida brasiliensis (Mexican free-tailed bat). The little and the big brown bats are common throughout the northern two-thirds of the country, while the Mexican free-tailed bat is the most common species in the southwest, sometimes even appearing in portions of the Southeast.
ANATOMY
WINGS
The finger bones of bats are much more flexible than those of other mammals, owing to their flattened cross-section and to low levels of minerals, such as calcium, near their tips. In 2006, Sears et al. published a study that traces the elongation of manual bat digits, a key feature required for wing development, to the upregulation of bone morphogenetic proteins (Bmps). During embryonic development, the gene controlling Bmp signaling, Bmp2, is subjected to increased expression in bat forelimbs - resulting in the extension of the offspring's manual digits. This crucial genetic alteration helps create the specialized limbs required for volant locomotion. Sears et al. (2006) also studied the relative proportion of bat forelimb digits from several extant species and compared these with a fossil of Lcaronycteris index, an early extinct species from approximately 50 million years ago. The study found no significant differences in relative digit proportion, suggesting that bat wing morphology has been conserved for over 50 million years.The wings of bats are much thinner and consist of more bones than the wings of birds, allowing bats to maneuver more accurately than the latter, and fly with more lift and less drag. By folding the wings in toward their bodies on the upstroke, they save 35 percent energy during flight. The membranes are also delicate, ripping easily; however, the tissue of the bat's membrane is able to regrow, such that small tears can heal quickly. The surface of their wings is equipped with touch-sensitive receptors on small bumps called Merkel cells, also found on human fingertips. These sensitive areas are different in bats, as each bump has a tiny hair in the center, making it even more sensitive and allowing the bat to detect and collect information about the air flowing over its wings, and to fly more efficiently by changing the shape of its wings in response. An additional kind of receptor cell is found in the wing membrane of species that use their wings to catch prey. This receptor cell is sensitive to the stretching of the membrane. The cells are concentrated in areas of the membrane where insects hit the wings when the bats capture them.
OTHER
The teeth of microbats resemble insectivorans. They are very sharp to bite through the hardened armor of insects or the skin of fruit.
Mammals have one-way valves in their veins to prevent the blood from flowing backwards, but bats also have one-way valves in their arteries.
The tube-lipped nectar bat (Anoura fistulata) has the longest tongue of any mammal relative to its body size. This is beneficial to them in terms of pollination and feeding. Their long, narrow tongues can reach deep into the long cup shape of some flowers. When the tongue retracts, it coils up inside its rib cage.
Bats possess highly adapted lung systems to cope with the pressures of powered-flight. Flight is an energetically taxing aerobic activity and requires large amounts of oxygen to be sustained. In bats, the relative alveolar surface area and pulmonary capillary blood volume are significantly larger than most other small quadrupedal mammals.
ECHOLOCATION
Bat echolocation is a perceptual system where ultrasonic sounds are emitted specifically to produce echoes. By comparing the outgoing pulse with the returning echoes, the brain and auditory nervous system can produce detailed images of the bat's surroundings. This allows bats to detect, localize, and even classify their prey in complete darkness. At 130 decibels in intensity, bat calls are some of the most intense, airborne animal sounds.
To clearly distinguish returning information, bats must be able to separate their calls from the echoes that they receive. Microbats use two distinct approaches.
Low duty cycle echolocation: Bats can separate their calls and returning echoes by time. Bats that use this approach time their short calls to finish before echoes return. This is important because these bats contract their middle ear muscles when emitting a call, so they can avoid deafening themselves. The time interval between the call and echo allows them to relax these muscles, so they can clearly hear the returning echo. The delay of the returning echoes provides the bat with the ability to estimate the range to their prey.
High duty cycle echolocation: Bats emit a continuous call and separate pulse and echo in frequency. The ears of these bats are sharply tuned to a specific frequency range. They emit calls outside of this range to avoid self-deafening. They then receive echoes back at the finely tuned frequency range by taking advantage of the Doppler shift of their motion in flight. The Doppler shift of the returning echoes yields information relating to the motion and location of the bat's prey. These bats must deal with changes in the Doppler shift due to changes in their flight speed. They have adapted to change their pulse emission frequency in relation to their flight speed so echoes still return in the optimal hearing range.
The new Yinpterochiroptera and Yangochiroptera classification of bats, supported by molecular evidence, suggests two possibilities for the evolution of echolocation. It may have been gained once in a common ancestor of all bats and was then subsequently lost in the Old World fruit bats, only to be regained in the horseshoe bats, or echolocation evolved independently in both the Yinpterochiroptera and Yangochiroptera lineages.
Two groups of moths exploit a bat sense to echolocate: tiger moths produce ultrasonic signals to warn the bats that they (the moths) are chemically protected or aposematic, other moth species produce signals to jam bat echolocation. Many moth species have a hearing organ called a tympanum, which responds to an incoming bat signal by causing the moth's flight muscles to twitch erratically, sending the moth into random evasive maneuvers.
In addition to echolocating prey, bat ears are sensitive to the fluttering of moth wings, the sounds produced by tymbalate insects, and the movement of ground-dwelling prey, such as centipedes, earwigs, etc. The complex geometry of ridges on the inner surface of bat ears helps to sharply focus not only echolocation signals, but also to passively listen for any other sound produced by the prey. These ridges can be regarded as the acoustic equivalent of a Fresnel lens, and may be seen in a large variety of unrelated animals, such as the aye-aye, lesser galago, bat-eared fox, mouse lemur, and others.
By repeated scanning, bats can mentally construct an accurate image of the environment in which they are moving and of their prey item.
OTHER SENSES
Although the eyes of most microbat species are small and poorly developed, leading to poor visual acuity, no species is blind. Microbats use vision to navigate, especially for long distances when beyond the range of echolocation, and species that are gleaners - that is, ones that attempt to swoop down from above to ambush tasty insects like crickets on the ground or moths up a tree - often have eyesight about as good as a rat's. Some species have been shown to be able to detect ultraviolet light, and most cave dwelling species have developed the ability to utilize very dim light. They also have high-quality senses of smell and hearing. Bats hunt at night, reducing competition with birds, minimizing contact with certain predators, and travel large distances (up to 800 km) in their search for food. Megabat species often have excellent eyesight as good as, if not better than, human vision; they need this for the warm climates they live in and the very social world they occupy, where relations and friends need to be distinguished from other bats in the colony. This eyesight is, unlike its microbat relations, adapted to both night and daylight vision and enables the bat to have some colour vision whereas the microbat sees in blurred shades of grey.
BEHAVIOUR
Most microbats are nocturnal and are active at twilight. A large portion of bats migrate hundreds of kilometres to winter hibernation dens, while some pass into torpor in cold weather, rousing and feeding when warm weather allows for insects to be active. Others retreat to caves for winter and hibernate for six months. Bats rarely fly in rain, as the rain interferes with their echolocation, and they are unable to locate their food.
The social structure of bats varies, with some leading solitary lives and others living in caves colonized by more than a million bats. The fission-fusion social structure is seen among several species of bats. The term "fusion" refers to a large numbers of bats that congregate in one roosting area, and "fission" refers to breaking up and the mixing of subgroups, with individual bats switching roosts with others and often ending up in different trees and with different roostmates.
Studies also show that bats make all kinds of sounds to communicate with others. Scientists in the field have listened to bats and have been able to associate certain sounds with certain behaviours that bats make after the sounds are made.
Insectivores make up 70% of bat species and locate their prey by means of echolocation. Of the remainder, most feed on fruits. Only three species sustain themselves with blood.
Some species even prey on vertebrates. The leaf-nosed bats (Phyllostomidae) of Central America and South America, and the two bulldog bat (Noctilionidae) species feed on fish. At least two species of bat are known to feed on other bats: the spectral bat, also known as the American false vampire bat, and the ghost bat of Australia. One species, the greater noctule bat, catches and eats small birds in the air.
Predators of bats include bat hawks, bat falcons and even spiders.
REPRODUCTION
Most bats have a breeding season, which is in the spring for species living in a temperate climate. Bats may have one to three litters in a season, depending on the species and on environmental conditions, such as the availability of food and roost sites. Females generally have one offspring at a time, which could be a result of the mother's need to fly to feed while pregnant. Female bats nurse their young until they are nearly adult size, because a young bat cannot forage on its own until its wings are fully developed.
Female bats use a variety of strategies to control the timing of pregnancy and the birth of young, to make delivery coincide with maximum food ability and other ecological factors. Females of some species have delayed fertilization, in which sperm is stored in the reproductive tract for several months after mating. In many such cases, mating occurs in the fall, and fertilization does not occur until the following spring. Other species exhibit delayed implantation, in which the egg is fertilized after mating, but remains free in the reproductive tract until external conditions become favorable for giving birth and caring for the offspring.
In yet another strategy, fertilization and implantation both occur, but development of the fetus is delayed until favorable conditions prevail, during the delayed development the mother still gives the fertilized egg nutrients, and oxygenated blood to keep it alive. However, this process can go for a long period of time, because of the advanced gas exchange system. All of these adaptations result in the pup being born during a time of high local production of fruit or insects.
At birth, the wings are too small to be used for flight. Young microbats become independent at the age of six to eight weeks, while megabats do not until they are four months old.
LIFE EXPECTANCY
A single bat can live over 20 years, but bat population growth is limited by the slow birth rate.
HUNTING, FEEDING AND DRINKING
Newborn bats rely on the milk from their mothers. When they are a few weeks old, bats are expected to fly and hunt on their own. It is up to them to find and catch their prey, along with satisfying their thirst.
HUNTING
Most bats are nocturnal creatures. Their daylight hours are spent grooming and sleeping; they hunt during the night. The means by which bats navigate while finding and catching their prey in the dark was unknown until the 1790s, when Lazzaro Spallanzani conducted a series of experiments on a group of blind bats. These bats were placed in a room in total darkness, with silk threads strung across the room. Even then, the bats were able to navigate their way through the room. Spallanzani concluded the bats were not using their eyes to fly through complete darkness, but something else.
Spallanzani decided the bats were able to catch and find their prey through the use of their ears. To prove this theory, Spallanzani plugged the ears of the bats in his experiment. To his pleasure, he found that the bats with plugged ears were not able to fly with the same amount of skill and precision as they were able to without their ears plugged. Unfortunately for Spallanzani, the twin concepts of sound waves and acoustics would not be understood for another century and he could not explain why specifically the bats were crashing into walls and the threads that he'd strung up around the room, and because of the methodology Spallanzani used, many of his test subjects died.
It was thus well known through the nineteenth century that the chiropteran ability to navigate had something to do with hearing, but how they accomplish this was not proven conclusively until the 1930s, by Donald R. Griffin, a biology student at Harvard University. Using a locally native species, the little brown bat, he discovered that bats use echolocation to locate and catch their prey. When bats fly, they produce a constant stream of high-pitched sounds. When the sound waves produced by these sounds hit an insect or other animal, the echoes bounce back to the bat, and guide them to the source.
FEEDING AND DIET
The majority of food consumed by bats includes insects, fruits and flower nectar, vertebrates and blood. Almost three-fourths of the world's bats are insect eaters. Bats consume both aerial and ground-dwelling insects. Each bat is typically able to consume one-third of its body weight in insects each night, and several hundred insects in a few hours. This means that a group of a thousand bats could eat four tons of insects each year. If bats were to become extinct, it has been calculated that the insect population would reach an alarmingly high number.
VITAMIN C
In a test of 34 bat species from six major families of bats, including major insect- and fruit-eating bat families, all were found to have lost the ability to synthesize vitamin C, and this loss may derive from a common bat ancestor, as a single mutation. However, recent results show that there are at least two species of bat, the frugivorous bat (Rousettus leschenaultii) and insectivorous bat (Hipposideros armiger), that have retained their ability to produce vitamin C. In fact, the whole Chiroptera are in the process of losing the ability to synthesize Vc which most of them have already lost.
AERIAL INSECTIVORES
Watching a bat catch and eat an insect is difficult. The action is so fast that all one sees is a bat rapidly change directions, and continue on its way. Scientist Frederick A. Webster discovered how bats catch their prey. In 1960, Webster developed a high-speed camera that was able to take one thousand pictures per second. These photos revealed the fast and precise way in which bats catch insects. Occasionally, a bat will catch an insect in mid-air with its mouth, and eat it in the air. However, more often than not, a bat will use its tail membrane or wings to scoop up the insect and trap it in a sort of "bug net". Then, the bat will take the insect back to its roost. There, the bat will proceed to eat said insect, often using its tail membrane as a kind of napkin, to prevent its meal from falling to the ground. One common insect prey is Helicoverpa zea, a moth that causes major agricultural damage.
FORAGE GLEANERS
These bats typically fly down and grasp their prey off the ground with their teeth, and take it to a nearby perch to eat it. Generally, these bats do not use echolocation to locate their prey. Instead, they rely on the sounds produced by the insects. Some make unique sounds, and almost all make some noise while moving through the environment.
FRUITS AND FLOWER NECTAR
Fruit eating, or frugivory, is a specific habit found in two families of bats. Megachiropterans and microchiropterans both include species of bat that feed on fruits. These bats feed on the juices of sweet fruits, and fulfill the needs of some seeds to be dispersed. The fruits preferred by most fruit-eating bats are fleshy and sweet, but not particularly strong smelling or colorful. To get the juice of these fruits, bats pull the fruit off the trees with their teeth, and fly back to their roosts with the fruit in their mouths. There, the bats will consume the fruit in a specific way. To do this, the bats crush open the fruit and eat the parts that satisfy their hunger. The remainder of the fruit, the seeds and pulp, are spat onto the ground. These seeds take root and begin to grow into new fruit trees. Over 150 types of plants depend on bats in order to reproduce.Some bats prefer the nectar of flowers to insects or other animals. These bats have evolved specifically for this purpose. For example, these bats possess long muzzles and long, extensible tongues covered in fine bristles that aid them in feeding on particular flowers and plants.[68] When they sip the nectar from these flowers, pollen gets stuck to their fur, and is dusted off when the bats take flight, thus pollinating the plants below them. The rainforest is said to be the most benefitted of all the biomes where bats live, because of the large variety of appealing plants. Because of their specific eating habits, nectar-feeding bats are more prone to extinction than any other type of bat. However, bats benefit from eating fruits and nectar just as much as from eating insects.
VERTEBRATES
A small group of carnivorous bats feed on other vertebrates and are considered the top carnivores of the bat world. These bats typically eat a variety of animals, but normally consume frogs, lizards, birds, and sometimes other bats. For example, one vertebrate predator, Trachops cirrhosus, is particularly skilled at catching frogs. These bats locate large groups of frogs by distinguishing their mating calls from other sounds around them. They follow the sounds to the source and pluck them from the surface of the water with their sharp canine teeth. Another example is the greater noctule bat, which is believed to catch birds on the wing.
Also, several species of bat feed on fish. These types of bats are found on almost all continents. They use echolocation to detect tiny ripples in the water's surface to locate fish. From there, the bats swoop down low, inches from the water, and use specially enlarged claws on their hind feet to grab the fish out of the water. The bats then take the fish to a feeding roost and consume the animal.
BLOOD
A few species of bats exclusively consume blood as their diet. This type of diet is referred to as hematophagy, and three species of bats exhibit this behavior. These species are the common, the white-winged, and the hairy-legged vampire bats. The common vampire bat typically consumes the blood of mammals, while the hairy-legged and white-winged vampires feed on the blood of birds. These species live only in Mexico, Central, and South America, with a presence also on the Island of Trinidad.
DEFECATION
Bat dung, or guano, is so rich in nutrients that it is mined from caves, bagged, and used by farmers to fertilize their crops. During the U.S. Civil War, guano was used to make gunpowder.
To survive hibernation months, some species build up large reserves of body fat, both as fuel and as insulation.
DRINKING
In 1960, Frederic A. Webster discovered bats' method of drinking water using a high-speed camera and flashgun that could take 1,000 photos per second. Webster's camera captured a bat skimming the surface of a body of water, and lowering its jaw to get just one drop of water. It then skimmed again to get a second drop of water, and so on, until it has had its fill. A bat's precision and control during flight is very fine, and it almost never misses. Other bats, such as the flying fox or fruit bat, gently skim the water's surface, then land nearby to lick water from chest fur.
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