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290:, appearing as pages in a book, are where gas exchange takes place. These appendages move rhythmically to drive blood in and out of the lamellae and to circulate water over them. Respiration being their main purpose, they can also be used for swimming in young individuals. If they are kept moist, the horseshoe crab can live on land for many hours.
286:) which have five pairs of them, the flap in front of them being the genital operculum which lacks gills. Book gills are flap-like appendages that effect gas exchange within water and seem to have their origin as modified legs. On the inside of each appendage, over 100 thin page-like membranes,
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Tetrapulmonata have two pairs of book lungs found on the second and third abdominal segments (Schizomida have lost a pair, and most advanced spiders have replaced at least one of the pairs with trachea). Scorpions have four pairs of book lungs, found on abdominal segments number three, four, five,
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One of the long-running controversies in arachnid evolution is whether the book lung evolved from book gills just once in a common arachnid ancestor, or whether book lungs evolved separately in several groups of arachnids as they came onto land. While the third abdominal segment in
Tetrapulmonata
269:. Although they have a similar book-like structure, book gills are external, while book lungs are internal. Both are considered appendages rather than conventional internal organs, as they develop from limb buds before the buds flatten into segmented
139:, have no traces of book lungs and breathe through tracheae or through their body-surfaces only. Gas exchange is performed by the thin walls inside the cavity instead, with their surface area increased by branching into the body as thin tubes called
31:
389:
Scholtz, Gerhard; Kamenz, Carsten (2006). "The book lungs of
Scorpiones and Tetrapulmonata (Chelicerata, Arachnida): Evidence for homology and a single terrestrialisation event of a common arachnid ancestor".
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Their number varies from just one pair in most spiders to four pairs in scorpions. The unfolded "pages" (plates) of the book lung are filled with hemolymph. The folds maximize the surface exposed to
74:. Each of these organs is located inside an open ventral abdominal, air-filled cavity (atrium) and connects with its surroundings through a small opening for the purpose of
143:. These tracheae may possibly have evolved directly from book lungs because the tracheae in some spiders have a small number of greatly elongated chambers.
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exchanged with the environment. In most species, no motion of the plates is needed to facilitate this kind of respiration.
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fossil lungs are almost indistinguishable from the lungs of modern arachnids, fully adapted to a terrestrial existence.
503:"Patterning mechanisms and morphological diversity of spider appendages and their importance for spider evolution"
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The pulmonate arachnids also appears to be the only members of
Arachnida where the respiratory pigment
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Pechmann, Matthias; Khadjeh, Sara; Sprenger, Frederik; Prpic, Nikola-Michael (November 2010).
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The origins of tetrapulmonate book lungs and their significance for chelicerate phylogeny
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have book lungs, the scorpions have a pair of sensory organs called pectines instead.
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Kamenz, Carsten; Dunlop, Jason A.; Scholtz, Gerhard; Kerp, Hans; Hass, Hagen (2008).
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The absence or presence of book lungs divides the
Arachnida into two main groups:
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In this spider diagram, the position of the book lungs is labelled 1.
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Internal anatomy of a female spider, book lungs shown in pink
104:. Stacks of alternating air pockets and tissue filled with
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used for atmospheric gas exchange that is present in many
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The oldest book lungs have been recovered from extinct
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The diversity and evolution of chelicerate hemocyanins
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Book gills are still present in the marine arthropod
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arachnids preserved in the 410 million-year-old
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give them an appearance similar to a "folded" book.
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261:Book lungs are thought to have evolved from book
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427:"Microanatomy of early Devonian book lungs"
265:, water-breathing structures among marine
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437:(2). London, UK: Royal Society: 212–215.
27:Type of lung commonly found in arachnids
474:Bhamrah, H. S.; Juneja, Kavita (2002).
339:. Oxford University Press US. pp.
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507:Arthropod Structure & Development
115:, and thereby maximize the amount of
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92:Book lungs are not related to the
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42:Spider book lungs (cross section)
558:Invertebrate respiratory system
480:. Anmol Publications Pvt. Ltd.
257:showing the legs and book gills
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477:An Introduction to Arthropoda
222:is present in their blood.
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404:10.1016/j.zool.2005.06.003
331:Foelix, Rainer F. (1996).
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519:10.1016/j.asd.2010.07.007
184:The a-pulmonate arachnids
96:of modern land-dwelling
155:The pulmonate arachnids
443:10.1098/rsbl.2007.0597
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82:Structure and function
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86:Further information:
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312:equivalent of blood.
158:book lungs present;
102:convergent evolution
237:of Scotland. These
189:microwhip scorpions
187:book lungs absent;
123:Occasionally absent
335:Biology of Spiders
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147:Arachnid taxonomy
60:respiration organ
16:(Redirected from
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398:(1): 2–13.
98:vertebrates
76:respiration
547:Categories
355:book lung.
319:References
245:Book gills
220:hemocyanin
209:sunspiders
193:harvestmen
168:Schizomida
137:harvestmen
131:, such as
66:, such as
18:Book lungs
532:20 August
310:arthropod
306:Hemolymph
294:Footnotes
215:and six.
205:Ricinulei
180:scorpions
172:Amblypygi
129:arachnids
106:hemolymph
68:scorpions
64:arachnids
56:book lung
527:20696272
461:18198139
412:16386884
288:lamellae
271:lamellae
239:Devonian
141:tracheae
452:2429929
392:Zoology
308:is the
279:Limulus
197:Acarina
176:spiders
72:spiders
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178:) and
174:, and
343:–64.
263:gills
133:mites
127:Many
94:lungs
534:2020
523:PMID
482:ISBN
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408:PMID
345:ISBN
135:and
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Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
