Deep-water Coral Reefs: Unique Biodiversity Hot-Spots (Springer Praxis Books)
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Toggle navigation Menu. Name of resource. Problem URL. Describe the connection issue. SearchWorks Catalog Stanford Libraries. Deep-water coral reefs : unique biodiversity hot-spots. Responsibility Martin Hovland. Physical description xxvi, p. Series Springer-Praxis books in life sciences incorporating aquatic and marine sciences.
Online Available online. Marine Biology Library Miller.
The four sample sites expose different facies. The samples collected directly west of below the Geomuseum Faxe consist of dense bryozoan-dominated rudstone with rare remains of corals, brachiopods, decapods, and bivalves site 3. It consists predominantly of branches and colony fragments of the mound-forming coral Dendrophyllia candelabrum , often only some centimeters apart, in between which some brachiopods, gastropods, bivalves, and decapods are found.
The octocoral packstone site site 1 , containing octocorals, scleractinians, and bryozoans primarily, is part of a dome-shaped structure surrounded by coral bafflestones. Other associated fossils are brachiopods, gastropods, bivalves, and decapods.
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These three sites are part of the type locality of the Faxe Formation [ 60 ]. The northernmost bafflestone site site 4 is composed of intermixed bryozoans and scleractinian corals, not as densely packed as the second site, but with many pockets filled with carbonate mud and abundant decapods as well as brachiopods, gastropods, and bivalves.
Dome-shaped octocoral site site 1 surrounded by scleractinian coral bafflestones. Sampling took place to the right of the small tree above the bucket blue at the level of the tree see also Additional file 1 : Figure S1. The site with the coral bafflestones site 2 is located towards the left of the dome not visible here. For counting the number of specimens per species, a number of standardizations were carried out to minimize biases see [ 29 ], for further discussion : 1 only internal molds with or without cuticle of carapaces with the axial part of the cervical groove preserved a unique landmark were counted; 2 specimens were collected by AAK only to minimize the possible effect of different sample strategies.
To investigate diversity differences among sites within the quarry, multiple measures of diversity were employed and all samples per site were merged for an adequate sample size for comparisons among sites. All specimens determined to the species-level were included in the analyses. These specimens were assigned to species based on the relative proportion of species of the respective taxon at each site to increase sample size Additional file 1 : Table S1.
Diversity per site was computed using: 1 species and genus richness, 2 individual rarefaction per site, and 3 the Shannon Index. Abundance data per species is unavailable for part of the localities, but they are for the Koskobilo and Faxe quarries so that diversity analyses 2 and 3 as above could be performed.
Deep-Water Coral Reefs: Unique Biodiversity Hot-Spots
All available specimens used here were systematically collected by AAK at both localities and samples were merged to increase sample size for all sites per locality. The maximum carapace width of decapods was measured for each possible specimen collected in Faxe measurements as in [ 29 ]. Width was chosen over length because width could be measured more easily for partial specimens due to the bilateral symmetry of decapods. The non-parametric Kruskal-Wallis test was used to compare widths among sites from the Faxe Quarry.
The Mann—Whitney pairwise comparisons test with and without a Bonferroni correction was used to assess whether pairs differed significantly. The same was done for the two most abundant species: Dromiopsis elegans [ 67 ] and D. Carapace widths and geometric means where possible of decapods and a subset true crabs or Brachyura, the most speciose decapod clade from Faxe were also compared, using the Mann—Whitney test, to those from the other localities with coral limestones as mentioned above.
For this purpose, the maximum carapace width and length were recorded as derived from the literature or as measured herein. Furthermore, the maximum widths of crab genera abundantly present in both Faxe and Koskobilo Caloxanthus and Faksecarcinus were also compared. Dromiopsis mosae [ 70 ] was recently suggested not to originate from the ENCI Quarry [ 71 ] and was not used. As Brachyura are the most speciose decapod clade known from the Paleocene, the maximum widths and geometric means of reef-associated crabs from Faxe were compared to those of all other Paleocene crabs.
Eye sockets or orbits are preserved for some decapod taxa and serve as a proxy for eye size. Eye socket height is measured here because socket width is frequently influenced by the eye stalks as well. As eye size is correlated with specimen size, carapace widths and lengths were also measured to obtain the geometric mean of the carapace. Differences in eye size between deep and shallow-water decapods are best expressed by comparing taxa within the same genus [ 72 ]. For that reason, eye socket heights of congeners from different inferred paleodepths are compared.
The epifaunal crabs of Caloxanthus were used for this purpose because they are abundant in Faxe Caloxanthus ornatus [ 48 ] and are also commonly found in shallow-water coral-associated rocks from the Albian of Koskobilo C. No other genera were suitable for this purpose. Specimens of museum collections were studied to obtain eye socket height and carapace size of Caloxanthus.
Using specimens of similar carapace size is important because eye size tends to scale with body size, but the performance of eyes e. Species that are almost always found at each site are Dromiopsis rugosus , D. Dromiopsis elegans is the most common species at each site, except for site 2 where D. These results were replicated when specimens not determined to the species-level were excluded.
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Carapaces of common decapod species from Faxe and one from Koskobilo. Carapaces from the Paleocene middle Danian of Faxe, Denmark a — f , and a carapace from the Albian of Koskobilo, Spain g — h , for comparison. Permission to use e — h [ 73 ] by The Palaeontological Association.
Scale bar width: 5. The slopes of the means middle lines suggest that site 4 exhibits the highest diversity. Rocks at all sites were lithified, but the carbonates at sites 1 and 3 were softer and more friable than other sites. This did not affect the recognition of decapods substantially, with the possible exception of site 1 where decapods were less well-preserved. All specimens used consist of complete or partial dorsal carapaces without venters and appendages attached.
Cuticle is preserved occasionally, but this did not influence the identification of specimens to the species-level here [ 74 ] because the taxonomy of decapods from Faxe is well-known [ 51 , 55 ]. The Shannon Index of Faxe 1. Both localities incorporate various microhabitats. Data for Koskobilo from [ 29 ]. After a Bonferroni correction, only the median sizes of sites 2 and 4 differ significantly using the Mann—Whitney test medians 7.
A larger size is also observed for the benthic crab Caloxanthus : maximum width for the species from Faxe is The single specimen of Caloxanthus known from the shallow-water, coral-associated Danian of Vigny [ 74 ] is also smaller 6. Conversely, the maximum width of the swimming crab Faksecarcinus appears larger in Koskobilo Although Dromiopsis praelaevior from the ENCI Quarry is only known from a single specimen [ 70 ], it exhibits a smaller maximum width compared to all four congenerics from Faxe Additional file 1 : Tables S3—7.
Likewise, specimens of Dromiopsis paucigranosa from Contrada Gecchelina di Monte di Malo and the Braggi Quarry [ 27 , 31 ] are smaller than Dromiopsis spp. Finally, the width and geometric means of Faxe crabs and all other Paleocene crabs Additional file 1 : Table S9 do not differ significantly Mann—Whitney U -values: , ; p -values: 0. The trend line of eye socket height versus maximum carapace width of C.
The eye socket height of the single specimen of C.
The hypothesis that decapod diversity is higher in coral facies compared to that of the intercalated bryozoan facies is partially supported. The coral-dominated facies site 2 harbors a higher diversity compared to the bryozoan-dominated facies site 3 Figs. This is consistent with the suggestion that decapod diversity in Eocene bryozoan-associated limestones from the USA South Carolina was lower than in coral-associated limestones in Europe, primarily due to ecological differences [ 75 ].
Although modern bryozoan habitats also contain a diverse, mostly facultative associated fauna including decapods [ 76 — 79 ], the relative diversity of this fauna compared that of coral reefs is unknown. At Faxe, decapod diversity and abundance is highest in the facies dominated by a mixture of bryozoans and scleractinian corals. This result cannot be explained by preservational differences because the coral-dominated site is composed of bafflestones of about equal hardness, whereas the bryozoan-dominated site is slightly softer, but decapod preservation is largely similar.
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The extensive, dense network of monotonous branching corals of primarily Dendrophyllia candelabrum at site 2 only some centimeters between each branch probably precluded inhabitation of an abundant, diverse decapod fauna, whereas a more varied, open framework with a higher number of microhabitats at site 4 attracted more decapods. This result may be consistent with data from modern cold-water coral habitats because: 1 the coral framework itself seems to harbor a relatively low diversity [ 7 , 35 , 36 ] and 2 habitat heterogeneity appears to promote diversity [ 34 , 69 , 80 ].
Preservation could have played a role for site 1, where sediments are much softer and decapod preservation is less pristine regardless of carapace size compared to other sites. Shrimps are most likely absent because they do not preserve well [ 81 , 82 ]. The relatively low decapod diversity of Faxe compared to that of other European, Cretaceous—Paleogene localities associated with corals Fig. This low diversity of decapods at Faxe is consistent with the relatively low number of reef-building coral species in modern cold-water reefs [ 12 ] and the relatively low fish species richness [ 4 ] compared to tropical reefs.
Moreover, the rarefaction trajectories also suggest a lower diversity for Faxe compared to Koskobilo Fig.
Seventy-nine percent of decapod families survived into the Paleogene, and many genera were able to survive in the sub tropical Americas, relatively close to the Chicxulub impact site [ 89 ]. Brachyurans also appear hardly affected [ 90 ]. Our new analyses of the uncorrected global standing decapod diversity show lower numbers of species and genera in the Danian than in the Maastrichtian. On the species-level, decapod and 75 brachyuran species from the Maastrichtian are known [ 30 ], whereas 92 decapod and 55 brachyuran species have been reported from the subsequent Danian e.
Seventy-five decapod genera including 51 brachyurans are reported from the Maastrichtian, whereas 55 decapod genera including 36 crabs are known from the Danian. However, when adjusted for the unequal duration of both stages and the differences in rock outcrop area Fig. Instead, Danian diversity is either approximately equal to that of the Maastrichtian or higher in the Danian.
Unlike for many other organisms worldwide [ 86 ], these results suggest a limited signal of a mass extinction in decapods and true crabs across this interval. Another factor that may have influenced the patterns is latitude because modern decapod diversity increases towards the equator [ 96 , 97 ]. The decapod carapaces at Faxe are generally well-preserved, also in comparison to, for example, Koskobilo pers.