Pteropoda

Pteropoda (common name pteropods, from the Greek meaning "wing-foot") are specialized free-
swimming pelagic sea snails and sea slugs, marine opisthobranch gastropods. Most live in top 10 m of Pteropoda
the ocean and are less than 1 cm long. The monophyly of Pteropoda is the subject of a lengthy debate; Temporal range:
they have even been considered as paraphyletic with respect to cephalopods.[1] Current consensus, Campanian–Recent
guided by molecular studies, leans towards interpreting the group as monophyletic.[2]
PreꞒ Ꞓ O S D C P T J K PgN
Pteropoda encompasses the two clades Thecosomata, the sea butterflies, and Gymnosomata, the sea
angels. The Thecosomata (lit. "case-body"[3]) have a shell, while the Gymnosomata ("naked body") do
not. The two clades may or may not be sister taxa; if not, their similarity (in that they are both pelagic,
small, and transparent, and both groups swim using wing-like flaps (parapodia) which protrude from
their bodies) may reflect adaptation to their particular lifestyle.

Contents A sea angel of the species Clione

Taxonomy limacina

Evolutionary history Scientific classification

Phylogeny Kingdom: Animalia
Vulnerability to ocean acidification Phylum: Mollusca
Distribution
Class: Gastropoda
References
Subclass: Heterobranchia
External links
Informal group: Opisthobranchia
Order: Pteropoda
Taxonomy Cuvier, 1804

The group Pteropoda was established by Georges Cuvier as "ptéropodes" in 1804.[4] François Péron
and Charles Alexandre Lesueur thought the group to be larger, and so they also included the opisthobranch taxa (Phyllirhoë and Glaucus), the
heteropoda taxa (Carinaria and Firola), and even the Ctenophora (Callianira). In 1810 these authors divided the whole group in two separate
groups: those with a shell and those without a shell.

In 1824 Henri Marie Ducrotay de Blainville named these two groups Gymnosomata and Thecosomata and named the combining order
Aporobranchia instead of Pteropoda.[5] He rejected the additional genera, except Phyllirhoë which he upgraded to a third group that he called
Psilosomata. Only much later was Phyllirhoë classified within the order Nudibranchia.

Other attempts were made to describe the Pteropoda. John Edward Gray divided the Pteropoda into Dactylobranchia (with just the genus
Cavolinia) and Pterobranchia (including all the other genera).[6] Cuvier (and his followers) did not accept the classification by de Blainville; they
preferred the original classification as described in Le Règne Animal.

In 1829 Paul Rang followed the Cuvierian classification, but tried to include the character of having a distinct head or not.[7] The German
naturalist Lorenz Oken went one step further and, for the sake of symmetry, wanted each order to contain four families and each family to contain
four genera.[8] Pierre André Latreille divided the Pteropoda according to the size of their fins: "Macroptérygiens" (including only
Pneumonoderma) and "Microptérygiens" (including all the others). In 1851 William Bullock Clark treated the Pteropoda as a family and emended
the spelling to Pteropodidae (a name now used for a family of fruit bats)

Finally all these attempts were abandoned and, as more and more species were described as a result of several scientific expeditions, the
classification of the Pteropoda into Thecosomata and Gymnosomata was generally adopted. Many of these new species were first described by
French zoologists, for example Jean René Constant Quoy and Joseph Paul Gaimard, Paul Rang, Alcide d'Orbigny and Louis François Auguste
Souleyet.

The relationship between these two clades is not unequivocally established, but it seems that they are sister taxa.[2]

Evolutionary history
Pteropods are estimated to have originated during the Early Cretaceous, around 133 million years ago, with the diversification into the major
lineages occurring during the mid-late Cretaceous. The oldest known fossil pteropod is a member of Limacinidae from the early-middle
Campanian deposits of the San Juan Islands.[9][10]

Phylogeny
Cladogram of genera and species of pteropods.
P d G
 Pteropoda Gymnosomata Clione antarctica

Pneumodermopsis spe

Spongiobranchaea australis

Pneumoderma violaceum

Cymbulia sibogae
Pseudothecosomata
Peracle reticulata
Euthecosomata Limacinoidea
Heliconoides inflatus

Limacina retroversa

Limacina antarctica

Limacina lesueurii

Limacina trochiformis

Limacina bulimoides

Creseis acicula

Creseis virgula
Thecosomata
Styliola subula

Hyalocylis striata

Clio pyramidata

Cuvierina atlantica
Cavolinioidea

Clio cuspidata

Diacavolinia
longirostris

Cavolinia inflexa

Diacria danae

Diacria trispinosa

The phylogenetic tree has been inferred from 2,654 nuclear proteins (representing 834,394 concatenated amino acid positions).[11]

Vulnerability to ocean acidification

A study was conducted on the West Coast of the United States to see ocean acidification’s effects on pteropods.[12] Limacina helicina was used to
test the sensitivity to decreasing pH.[12] This species of pteropod is potentially vulnerable to the corrosive waters associated with ocean
acidification due to their calcium carbonate shell.[13] The shell of a pteropod was immersed in ocean water with the projected pH level that the
water will reach by the year 2100. After a month and a half in the water, the shell had almost completely dissolved.[12]
Distribution
Pteropods are found in all major oceans, usually 0–10 metres (0–33 ft) below the ocean surface and in all
levels of latitude. Pteropods can be found lower than 10 meters, but in less amounts in terms of biomass,
however pteropod distribution is more spread out deeper based on findings. This can be explained as
Pteropods from tropical areas become more common in deeper areas. They are not found commonly in the
deep sea, in fact, few live lower than 500 meters below sea level. Continental shelves, areas containing
many opportunities for nutrients, and productivity are locations in which Pteropods are usually populous,
according to patterns in data. Springtime is a peak season for pteropoda, as they reach higher populations,
though data shows that pteropoda south of the equator are less abundant seasonally. In addition, current
data suggests that, 93% of the world's pteropods are part of the Thecosomata family, while the 7% are Unhealthy pteropod showing effects
Gymnosomata.[14] of ocean acidification

References
1. Wägele, Heike; Klussmann-Kolb, Annette; Verbeek, Eva; Schrödl, Michael (2013). "Flashback and foreshadowing—a review of
the taxon Opisthobranchia" (https://doi.org/10.1007%2Fs13127-013-0151-5). Organisms Diversity & Evolution. 14: 133–149.
doi:10.1007/s13127-013-0151-5 (https://doi.org/10.1007%2Fs13127-013-0151-5).
2. Klussmann-Kolb, A.; Dinapoli, A. (2006). "Systematic position of the pelagic Thecosomata and Gymnosomata within
Opisthobranchia (Mollusca, Gastropoda) - revival of the Pteropoda". Journal of Zoological Systematics and Evolutionary
Research. 44 (2): 118. doi:10.1111/j.1439-0469.2006.00351.x (https://doi.org/10.1111%2Fj.1439-0469.2006.00351.x).
3. "theco-" (https://oed.com/search?searchType=dictionary&q=theco-). Oxford English Dictionary (Online ed.). Oxford University
Press. (Subscription or participating institution membership (https://www.oed.com/public/login/loggingin#withyourlibrary) required.)
4. Mémoire sur l'Hyale et Ie Pneumoderme; Ann. Mus. Hist. Nat. Paris., 4 p. 232)
5. Diet. d. Sci. Nat., t. xxxii. p. 271.
6. London Medical Repository, p. 235, 1821.
7. Manuel de l’histoire naturelle des mollusques et leurs coquilles
8. Description d'un genre nouveau de la classe des Ptéropodes, Ann. d. &i. Nat., ser. 1, t. V. p. 284, 1825.
9. Peijnenburg, Katja T. C. A.; Janssen, Arie W.; Wall-Palmer, Deborah; Goetze, Erica; Maas, Amy E.; Todd, Jonathan A.; Marlétaz,
Ferdinand (2020-10-13). "The origin and diversification of pteropods precede past perturbations in the Earth's carbon cycle" (htt
ps://www.ncbi.nlm.nih.gov/pmc/articles/PMC7568333). Proceedings of the National Academy of Sciences. 117 (41): 25609–
25617. doi:10.1073/pnas.1920918117 (https://doi.org/10.1073%2Fpnas.1920918117). PMC 7568333 (https://www.ncbi.nlm.nih.
gov/pmc/articles/PMC7568333). PMID 32973093 (https://pubmed.ncbi.nlm.nih.gov/32973093).
10. A. W. Janssen, J. L. Goedert, Notes on the systematics, morphology and biostratigraphy of fossil holoplanktonic Mollusca, 24.
First observation of a genuinely Late Mesozoic thecosomatous pteropod. Basteria 80, 59–63 (2016)
11. Peijnenburg, Katja T. C. A.; Janssen, Arie W.; Wall-Palmer, Deborah; Goetze, Erica; Maas, Amy E.; Todd, Jonathan A.; Marlétaz,
Ferdinand (2020-09-24). "The origin and diversification of pteropods precede past perturbations in the Earth's carbon cycle" (htt
ps://www.ncbi.nlm.nih.gov/pmc/articles/PMC7568333). Proceedings of the National Academy of Sciences. 117 (41): 25609–
25617. doi:10.1073/pnas.1920918117 (https://doi.org/10.1073%2Fpnas.1920918117). ISSN 0027-8424 (https://www.worldcat.o
rg/issn/0027-8424). PMC 7568333 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7568333). PMID 32973093 (https://pubmed.
ncbi.nlm.nih.gov/32973093).
12. Bednaršek, N.; Feely, R. A.; Reum, J. C. P.; Peterson, B.; Menkel, J.; Alin, S. R.; Hales, B. (2014). "Limacina helicina shell
dissolution as an indicator of declining habitat suitability owing to ocean acidification in the California Current Ecosystem" (http
s://www.ncbi.nlm.nih.gov/pmc/articles/PMC4024287). Proc. R. Soc. B. 281 (1785): 20140123. doi:10.1098/rspb.2014.0123 (http
s://doi.org/10.1098%2Frspb.2014.0123). ISSN 0962-8452 (https://www.worldcat.org/issn/0962-8452). PMC 4024287 (https://ww
w.ncbi.nlm.nih.gov/pmc/articles/PMC4024287). PMID 24789895 (https://pubmed.ncbi.nlm.nih.gov/24789895).
13. Comeau, S.; Gorsky, G.; Jeffree, R.; Teyssié, J.-L.; Gattuso, J.-P. (2009-09-04). "Impact of ocean acidification on a key Arctic
pelagic mollusc (Limacina helicina)" (https://doi.org/10.5194%2Fbg-6-1877-2009). Biogeosciences. 6 (9): 1877–1882.
doi:10.5194/bg-6-1877-2009 (https://doi.org/10.5194%2Fbg-6-1877-2009). ISSN 1726-4189 (https://www.worldcat.org/issn/172
6-4189).
14. N. Bednaršek, J. Možina, M. Vogt, C. O'Brien, and G. A. Tarling (10 Dec 2012). "The global distribution of pteropods and their
contribution to carbonate and carbon biomass in the modern ocean" (https://www.earth-syst-sci-data.net/4/167/2012/). Earth
System Science Data. 4 (1): 167–186. doi:10.5194/essd-4-167-2012 (https://doi.org/10.5194%2Fessd-4-167-2012).
hdl:20.500.11850/60379 (https://hdl.handle.net/20.500.11850%2F60379). Retrieved 18 June 2018.

External links
Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition auf dem Dampfer "Valdivia" 1898-1899. Part 9 Atlas (https://w
ww.biodiversitylibrary.org/item/29027) by German planktologist Carl Chun
Plankton Chronicles (https://web.archive.org/web/20120112073516/http://www.planktonchronicles.org/en/episode/17) Short
documentary films & photos
http://www.marinespecies.org/aphia.php?p=taxdetails&id=325345
Report of the Scientific results of the Voyage of the H.M.S. Challenger (1873-1876), Zoology, part LVIII (1887) : Report on the
Pteropoda by Paul Pelseneer (https://web.archive.org/web/20120306073931/http://www.19thcenturyscience.org/HMSC/HMSC-
Reports/Zool-58/README.htm)
Alice K. Burridge, Christine Hörnlein, Arie W. Janssen, Martin Hughes, Stephanie L. Bush, Ferdinand Marlétaz, Rebeca Gasca,
Annelies C. Pierrot-Bults, Ellinor Michel, Jonathan A. Todd, Jeremy R. Young, Karen J. Osborn, Steph B. J. Menken, Katja T. C.
A. Peijnenburg : Time-calibrated molecular phylogeny of pteropods; PLOS|ONE, June 12, 2017 (https://journals.plos.org/ploson
e/article?id=10.1371/journal.pone.0177325)
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