Mostrando entradas con la etiqueta Astrotoma. Mostrar todas las entradas
Mostrando entradas con la etiqueta Astrotoma. Mostrar todas las entradas

Gigantism and dwarfism


Gigantism is well known among Antarctic benthic organisms, although the fact is that size abnormalities occur in both directions (gigantism and dwarfism). Is it possible that it would be a case of evolution with a disruptive selection in which the organisms with extreme phenotypes would be the dominant forms?  That means, being small size organisms (ophiuroids with disc less than 5mm) and large size ones (ophiuroids with disk of about 30mm) more abundant in species compared to the middle size organisms (ophiuroids with disk of approximately 10 mm).



The origin of dwarfism seems to be found in the difficulty to precipitate calcium carbonate at low temperatures, which is supposed to be a limiting factor for organisms with calcareous skeletons, such as calcareous foraminifera, prosobranch gastropods, bivalves and brachiopods, among which we can find Antarctic species of very small size. Antarctic echinoderms with a skeleton composed of calcareous dermal ossicles are “doomed” to higher energy consumption in order to incorporate calcium carbonate, compared to echinoderms from other geographical areas, for this reason, it seems logical to consider this enviroment to be a hostile one for this zoological group development, although it does not look to be like that, as  the echinoderms are among the benthic groups with most diversity and biomass in the Antarctic benthos.
Whatever the case and considering dwarf ophiuroid species those that do not reach 5 mm disk diameter , we have: Ophiacantha paramedea, Ophiomitrella ingrata, Amphiophiura antarctica, Ophiocten banzarei, Ophiocten bisquamatum, Ophiomastus conveniens, Ophiomastus ludwigi, Ophiomastus perforatus, Ophiomastus primula, Ophiomastus trispinosus, Ophiosteira bullivanti, Ophiopyrgus australis, Amphiura lymani,  Amphiura algida, Amphiura microplax, Amphiura monorima and Ophiozonella antarctica.


Antarctic animals with siliceous skeletons or those with no mineralized skeleton, can multiply by up to ten the size of the individuals of the species compared to ones of the same groups in other geographical areas, as it happens to nemerteans, pycnogonids and giant isopods, like if meeting the Bergmann’s rule. The waters with low temperatures and seasonal food shortages (in winter) reduce basal metabolism which may cause a delay in sexual maturity and increased longevity, the two factors that, along with  an environment with little amount of predators can favor the development of a giant size.
Among the ophiuroids, even despite its calcareous skeleton, appear giant Antarctic species, considering giant the species with disk diameter over 30 mm, we have: Gorgonocephalus chilensis  (though the gorgonocephalids in general have large size) Astrotoma agassizii, Ophiosparte gigas, Ophionotus victoriae, Ophionotus hexactis, Ophiura flexibilis, Ophiura lenticularis, Ophiocamax gigas and Ophiocamax drygalskii.

Taking the data of 117 species of ophiuroids present in Antarctic waters (I’ve excluded those with high distribution outside Antarctic waters) and represented according to disk sizes in ranges, we obtain the following graphic:


It shows a classic Gaussian bell curve, which clearly indicates an evolutionary “stabilizing type” tendency, i.e. where corporal phenotypes of middle size have been selected predominantly
When compared with species from other latitudes as in the case of brittle stars found in British waters (excluding cosmopolitan species), we have the following graphic as a result:


Where we can see presented proportionally even more anomalous cases respecting size.
Therefore it doesn’t seem to be that the dwarfism or gigantism have been selective factors at group level in the Antarctic ophiuroids, but it is more about cases that have to be treated in a particular way.

References
ARNAUD, P.M. 1974. Contribution a la bionomie marine benthique des regions antarctiques et subantartiques. Téthys, 6, 467–653.

SOUTHWARD, E.C. & CAMPBELL, A.C. 2006. Echinoderms. Synopses of the British Fauna. Edited by Crothers, J.H. & Hayward, P.J. The Linnean Society of London and Estuarine and Coastal Sciences Association.

TORTONESE, E. 1965. Echinodermata. Fauna D’Italia. Edizioni Calderini, Bologna.

November 2010

Some peculiarities of the Antarctic ophiuroids

Ophiozonella sp. and Ophioceres sp. in ventral view

Mixture of Paleozoic and deep sea fauna
The benthos of shallow-waters in Antarctica is a mixture of:
–       Faunas that are common to Australia and New Zealand,  South America, South Africa, but also with a large number of endemics.
–       Similarities to paleozoic faunas.
–       Invasion of the organisms proceeding from the adjacent deep-sea communities.

The absence of crabs and sharks, as well as scarce biodiversity and biomass of teleosts implies a low predation on exoskeleton zoological groups, which causes an abundance in biodiversity and biomass of echinoderms (Aroson & Blake 2001).
Viviparism
Among the Antarctic ophiuroids there is a higher proportion of viviparous species than in other marine areas. Such a thing does not seem to be an adaptation to low temperatures, but to the geographic type of insular habitats of shallow waters and to the insulation by the circumpolar current, the adaptation that prevents the dispersal of larvae Ostergren (1912).
There are viviparous species that are multibrachial (like Ophiacantha vivipara and Ophionotus hexactis), and there are species which are proximate to the aforementioned ones that, being pentamers, are not viviparous (like Ophiacantha pentactis and Ophionotus victoriae). Such a mechanism permits to accommodate more embryos by having more bursae Smirnov (1984).

Ophionotus hexactis with an embryo
Incubation period
The viviparous species have long incubation period (in comparison with the tropical species or the species from temperate waters) and very slow rhythm of growth, measured with Von Bertalanffy growth constant, the fact that indicates that large size individuals are very long-lived, like, for example, in the case of Astrotoma agasiizii, which is one of the largest ophiuroids and whose age calculated from vertebral growth rings can reach 91 years Dahm (1996).

Gigantism
The cases of gigantism among some representatives of the Antarctic fauna can also be found among the ophiuroids; so, while the average disk size of ophiuroids, generally measured in millimeters, is about 10-15 mm, there are some Antarctic ophiuroids with disk size of about some centimeters like, for example, Ophionotus victoriae with disk diameter up to 43 mm, Ophiosparte gigas, up to 60 mm, or Astrotoma agasiizii, up to 60 mm.
Vertebra with growth rings, Ophionothus victoriae
A,distal surface; B,proximal surface; 1,canal for radial nerve; 2,fossa for lower intervertebral muscle; 3,fossa for upper intervertebral muscles; 4, central projection; 5, central depression

Vertebra with growth rings, Astrotoma agassizii
A,distal surface; B,proximal surface  1, hour-glass projection; 2, canal for radial nerve

Can they adapt?
Another peculiarity that suggest  the investigations of Peck et al (2009) may be the limited adaptability to environmental variations. His research in aquarium with increasing temperature give for Ophionotus victoriae a scarce survival time. The ophiuroids begin to die from the 24th day at 2 º C, and the 19th at 3 ° C. It seems to suggest that the current global climate change could cause the loss of biodiversity in the Antarctic benthos because of a small temperature increase.

References
Aronson RB, Blake D (2001) Global Climate Change and the Origin of Modern Benthic Communities in Antarctica AMER. ZOOL., 41:27–39

Dahm (1996) Ecology and Population Dynamics of Antarctic Ophiuroids (Echinodermata). Ber. Polarforsch. 194

Peck LL S, Massey A, Thorne M A S, Clark M S (2009) Lack of acclimation in Ophionotus victoriae: brittle stars are not fish. Polar Biol 32: 399-402

Peck LL S,  Antartctic marine stenotherms, temperature seasonality and change. Powerpoint presentation.

Östergren, H (1912) Über dieBrutpfl ege der Echinodermen in den Südpolaren Küstengenbieten. Zeitschrift Wissenschaften Zoologie, 101:325– 341

Smirnov IS (1984)  Fauna of antarctic and subantarctic brittle-stars. Ph.D . Leningrad. 1984

January 2010

Astrotoma agassizii Lyman, 1875


One of the most emblematic animals from Antarctic waters is the euryalid Astrotoma agassizii Lyman, 1875. Gigant, long-lived, with a disk diameter of up to 70 mm and an age which could overpass 100 years; Dahm (1996) calculated, by the growth rings of the arm vertebrae, the age of 90 years for a specimen of disc less than 50 mm.

Astrotoma is a predator, who climbs up the gorgonians from Primnoella and Thouarella genus, waits patiently for the arrival of copepods which will be caught by its ambulacral tentacles Dearborn et al (1986). His rough and tough skin defends it from the nematocysts that the polyps of the Gorgon throw without success. Astrotoma grabs the gorgonia with its modified arm spines and the hooklets that cover the skin of its arms arranged in rings.

1. ambulacral pore, 2. arm spine, 3. arm ring with hooklets, 4. jaws, 5. genital slit, 6. radial shields, 7. madreporite

Astrotoma is a hermaphrodite incubator Bernasconi (1965), something common in Antarctic Brittle stars, but the thing that is not so common is that, without a planktonic development that would help to expand its geographical distribution, it however presents a wide circumpolar distribution in shelf waters, in addition to being presented in the waters of Magellan. The Antarctic Circumpolar Current is a divisor of polar and magellanic populations Hunter (2008), there is no gene flow but these are isolated populations, molecularly distinct, although it doesn’t seem to be any morphological differentiation. Smirnov (1986) suggested the possibility of phoresis by macrophytes, an interesting option that could explain their wide distribution.

Arm ring with hooklets

References

Bernasconi I (1965) Astrotoma agassizii Lyman, especie vivipara del Atlántico Sur. Physis 25 (69): 1-5

Dahm C (1996) Ökologie und populationsdynamik antarktischer ophiuroiden (Echinodermata). Ber Polarforsch 194: 1-289

Dearborn, JH, Ferrari FD,  Edwards KC (1986) Can pelagic aggregations cause benthic satiation? Feeding biology of the Antarctic brittle star Astrotoma agassizii (Echinodermata: Ophiuroidea). Ant. Res. Ser., 44 (Biology of the Antarctic Seas XVII): 1-28

Hunter RL & Halanych KM  (2008) Evaluating Connectivity in the Brooding Brittle Star Astrotoma agassizii across the Drake Passage in the Southern Ocean. J Hered 99:137-148

Smirnov IS (1984)  Fauna of antarctic and subantarctic brittle-stars. Ph.D . Leningrad. 1984 [in Russian]

Collected by Pablo J. González-López. Cruiser ANTARKTIS XXIII/8  Polarstern

May 2010