Book lung

A book lung is a type of respiration organ used for atmospheric

gas exchange that is found in many arachnids, such as scorpions
and spiders. Each of these organs is found inside an open ventral
abdominal, air-filled cavity (atrium) and connects with the
surroundings through a small opening for the purpose of
respiration.

Contents
In this spider diagram, the book lung
Structure and function is labelled 1.
Occasionally absent
Arachnid taxonomy
Book gills
Footnote
References

Structure and function Spider book lungs (cross section)

Book lungs are not related to the lungs of modern land-dwelling

vertebrates. Their name describes their structure. Stacks of
alternating air pockets and tissue filled with hemolymph[a] give
them an appearance similar to a "folded" book.[1]

Their number varies from just one pair in most spiders to four
pairs in scorpions. The unfolded "pages" (plates) of the book lung
Book lungs of spider (shown in pink)
are filled with hemolymph. The folds maximize the surface
exposed to air, and thereby maximize the amount of gas
exchanged with the environment. In most species, no motion of the plates is needed to facilitate this kind
of respiration.

Occasionally absent
Sometimes, book lungs can be absent, and 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 tracheae. These
tracheae may possibly have evolved directly from the book lungs because the tracheae in some spiders
have a small number of greatly elongated chambers. Many arachnids, such as mites and harvestmen
(Opiliones), have no traces of book lungs and breathe through tracheae or through their body surfaces
only.

Arachnid taxonomy
The absence or presence of book lungs divides the Arachnida into two main groups:
The pulmonate arachnids
book lungs present; scorpions and the Tetrapulmonata; whip scorpions, Schizomida,
Amblypygi, and spiders
The a-pulmonate arachnids
book lungs absent; microwhip scorpions, harvestmen, Acarina, pseudoscorpions,
Ricinulei and sunspiders

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,[2] or whether book lungs evolved separately in several
groups of arachnids as they came onto land.

The oldest book lungs have been recovered from extinct trigonotarbid arachnids preserved in the
410 million-year-old Rhynie chert of Scotland. These Devonian fossil lungs are almost indistinguishable
from the lungs of modern arachnids, fully adapted to a terrestrial existence.[3]

Book gills
It is believed that book lungs evolved from book gills. Although
they have a similar book-like structure, book gills are found
externally, while book lungs are found internally.[4]

Book gills are still found in the marine arthropod Limulus

(horseshoe crabs) 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, lamellae, Underside of a female horseshoe
crab showing the legs and book gills
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.

Horseshoe crabs' book gills are developed from the base of the abdominal or opisthosomal appendages.
These five pairs of appendages are flap-like and membranous, with the under-surface of each flap formed
into many leaf-like folds called lamellae. Thus each gill bears hundreds of thin membranous lamellae
arranged like pages in a book. Each lamella is provided with capillaries separated from the outer sea
water by a thin wall that acts as the osmoregulatory membrane.The movement of the organs maintain a
constant water flow for gaseous exchange. The oxygen is diffused in from the water and the carbon
dioxide is expelled out. The coxae of the last pair of appendages bear short stapulate flagella which helps
to clean the gills and serves to sense the oxygen content of the water current.

Footnote
a. Hemolymph is the arthropod equivalent of blood.

References
1. Foelix, Rainer F. (1996). Biology of Spiders (https://books.google.com/books?id=XUgyqxNK
hyAC&pg=PA64&dq=book+lung#PPA64,M1). Oxford University Press US. pp. 61–64.
 ISBN 0-19-509594-4.
2. 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". Zoology. 109: 2–13.
doi:10.1016/j.zool.2005.06.003 (https://doi.org/10.1016%2Fj.zool.2005.06.003).
3. Kamenz, Carsten; Dunlop, Jason A.; Scholtz, Gerhard; Kerp, Hans; Hass, Hagen (2008).
"Microanatomy of early Devonian book lungs" (http://rsbl.royalsocietypublishing.org/content/
4/2/212). Biology Letters. London, UK: Royal Society. 4: 212–215.
doi:10.1098/rsbl.2007.0597 (https://doi.org/10.1098%2Frsbl.2007.0597).
4. Bhamrah, H. S.; Juneja, Kavita (2002). An Introduction to Arthropoda (https://books.google.
com/books?id=b_gsiQXNhTUC&pg=PA316&dq=book+lung+book+gills). Anmol Publications
Pvt. Ltd. ISBN 81-261-0673-5.

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