Bryozoans as palaeoenvironmental indicators. Paul D Taylor, NHM, London
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1 Bryozoans as palaeoenvironmental indicators Paul D Taylor, NHM, London
2 What are bryozoans?
3 What are bryozoans? invertebrate phylum
4 What are bryozoans? invertebrate phylum >6000 living species
5 What are bryozoans? invertebrate phylum >6000 living species mainly marine
6 What are bryozoans? invertebrate phylum >6000 living species mainly marine intertidal to 8000 m depth
7 What are bryozoans? invertebrate phylum >6000 living species mainly marine intertidal to 8000 m depth suspension feeders
8 What are bryozoans? invertebrate phylum >6000 living species mainly marine intertidal to 8000 m depth suspension feeders all colonial
9 What are bryozoans? invertebrate phylum >6000 living species mainly marine intertidal to 8000 m depth suspension feeders all colonial usually sessile, benthic
10 What are bryozoans? invertebrate phylum >6000 living species mainly marine intertidal to 8000 m depth suspension feeders all colonial usually sessile, benthic most have CaCO 3 skeletons
11 What are bryozoans? invertebrate phylum >6000 living species mainly marine intertidal to 8000 m depth suspension feeders all colonial usually sessile, benthic most have CaCO 3 skeletons Ordovician-Recent
12 Bryozoan colony-forms
13 Robert Hooke s illustration of Flustra from Micrographia (1665)
14
15
16 125 Phanerozoic bryozoan family diversity 100 families cheilostomes ORDOVICIAN Ma PRESENT DAY
17 125 Phanerozoic bryozoan family diversity 100 families Ordovician radiation cheilostomes ORDOVICIAN Ma PRESENT DAY
18 125 Phanerozoic bryozoan family diversity 100 families Ordovician radiation Permian extinction cheilostomes ORDOVICIAN Ma PRESENT DAY
19 125 Phanerozoic bryozoan family diversity 100 families Ordovician radiation Permian extinction Cretaceous radiation cheilostomes ORDOVICIAN Ma PRESENT DAY
20 Abundant Upper Ordovician Cincinnati, USA rock-forming
21 Abundant Diverse Upper Ordovician Cincinnati, USA rock-forming Upper Cretaceous Maastricht, Netherlands > 600 bryozoan species
22 Bryozoans as palaeoenvironmental indicators 1. modern environments with bryozoans 2. colony-form analysis 3. branch diameter as a depth indicator 4. zooid size as a temperature indicator 5. palaeolatitudinal distribution of bryozoan carbonates through time [skeletal geochemistry] Akkeshi Bay Japan
23 1. modern environments with bryozoans
24 Modern environments colonized by bryozoans
25 Modern environments colonized by bryozoans Freshwater bryozoans Low diversity Unmineralized
26 Modern environments colonized by bryozoans Freshwater bryozoans Low diversity Unmineralized Brackish bryozoans Low diversity Weakly mineralized
27 Modern environments colonized by bryozoans Freshwater bryozoans Low diversity Unmineralized Brackish bryozoans Low diversity Weakly mineralized Deep sea bryozoans Moderate diversity Weakly mineralized
28 Modern environments colonized by bryozoans Freshwater bryozoans Low diversity Unmineralized Brackish bryozoans Low diversity Weakly mineralized Shelf bryozoans High diversity Most species well-mineralized Deep sea bryozoans Moderate diversity Weakly mineralized
29 Recent bryozoan species diversity increasing with depth English Channel (Grant & Hayward 1985)
30 Bryozoan assemblage species richness vs depth in the eastern (black dots) and western (white dots) North Atlantic 10 m 75 m Clarke & Lidgard Journal of Animal Ecology 69:
31 2. colony-form analysis
32 Encrus'ng: Porella
33 Encrus'ng: Porella Dome shaped: 'Dianulites'
34 Encrus'ng: Porella Dome shaped: 'Dianulites' Narrow branched: Arcanopora
35 Encrus'ng: Porella Dome shaped: 'Dianulites' Robust branched: Heteropora Narrow branched: Arcanopora
36 Encrus'ng: Porella Dome shaped: 'Dianulites' Palmate: Phaenopora Robust branched: Heteropora Narrow branched: Arcanopora
37 Encrus'ng: Porella Dome shaped: 'Dianulites' Foliose: Pentapora Palmate: Phaenopora Robust branched: Heteropora Narrow branched: Arcanopora
38 Dome shaped: 'Dianulites' Encrus'ng: Porella Foliose: Pentapora Fenestrate: Chasmatopora Palmate: Phaenopora Robust branched: Heteropora Narrow branched: Arcanopora
39 Dome shaped: 'Dianulites' Encrus'ng: Porella Foliose: Pentapora Fenestrate: Chasmatopora Ar'culated: Cellaria Palmate: Phaenopora Robust branched: Heteropora Narrow branched: Arcanopora
40 Dome shaped: 'Dianulites' Encrus'ng: Porella Foliose: Pentapora Fenestrate: Chasmatopora Ar'culated: Cellaria Free- living: Cupuladria Palmate: Phaenopora Robust branched: Heteropora Narrow branched: Arcanopora
41 Cheilostome bryozoan colony-forms (from Moissette 2000) can these colony-forms be used as environmental indicators?
42 Cheilostome bryozoan colony-forms (from Moissette 2000) deep water shallow water strong currents high sedimentation can these colony-forms be used as environmental indicators?
43 Sea-bed topography around New Zealand Otago Shelf
44 bryozoan meadow, 100 metres Otago Shelf, New Zealand sediment from bryozoan meadow
45 Different colony-forms represent alternative functional adaptive strategies Encrus'ng colonies Uniserial runner Multiserial sheet
46 37 zooid encrusting colonies on substrates divided into equal-sized squares sheet runner
47 37 zooid encrusting colonies on substrates divided into equal-sized squares sheet runner Runner samples more squares of substrate and spreads a greater distance from origin (fugitive strategy)
48 37 zooid encrusting colonies on substrates divided into equal-sized squares sheet runner Runner samples more squares of substrate and spreads a greater distance from origin (fugitive strategy) Sheet is compact and better able to defend space ( confrontational strategy )
49 Percentage of colony-forms in bryozoan assemblages from shallow and deep stations in Spitsbergen SHALLOW DEEP (> 50 m) 14% 21% 74% Multiserial encrusters Uniserial encrusters 77%?reflecting decreasing food resources with depth
50 Fenestrate colonies (reteporiform) Cheilostome 'Sertella' (Oligocene, Argen'na) 'traditionally' regarded as adaptation to strong currents but colonies can be found in slow flow regimes too (e.g. caves) Chasmatopora (Ordovician, Estonia)
51 Ar'culated colonies (cellariiform) Cheilostome Cellaria (Recent, Croa'a) elastic nodes allow flexure in strong currents Crisia (Recent, Brazil) but colonies also found in areas of relatively high sedimentation and weak currents sediment can be shaken off branches
52 Free living colonies (lunuli'form) Cupuladria (Miocene, France) Cupuladria (Pliocene, England) able to live on fine sediments (fine sand, silt and mud) where no large substrates are present colonies can dig themselves out if buried
53 adapted for living on fine-grained, unstable substrates
54 Dome shaped: 'Dianulites' Encrus'ng: Porella Foliose: Pentapora Fenestrate: Chasmatopora Ar'culated: Cellaria Free- living: Cupuladria Palmate: Phaenopora Robust branched: Heteropora Narrow branched: Arcanopora
55 Changing proportion of bryozoan colony-forms through time ORD SIL DEV CARB PERM TRI JUR CRET PG NG
56 Using bryozoan colony-form in palaeoenvironmental reconstruction must be approached with caution
57 Using bryozoan colony-form in palaeoenvironmental reconstruction must be approached with caution particular colony-forms are seldom if ever restricted to one environment; sometimes almost all major colony-forms can be found together
58 Using bryozoan colony-form in palaeoenvironmental reconstruction must be approached with caution particular colony-forms are seldom if ever restricted to one environment; sometimes almost all major colony-forms can be found together not enough known about factors controlling colony-forms in modern environments
59 Using bryozoan colony-form in palaeoenvironmental reconstruction must be approached with caution particular colony-forms are seldom if ever restricted to one environment; sometimes almost all major colony-forms can be found together not enough known about factors controlling colony-forms in modern environments colony-forms represent alternative functional adaptive strategies that may be viable in many different environments
60 Using bryozoan colony-form in palaeoenvironmental reconstruction must be approached with caution particular colony-forms are seldom if ever restricted to one environment; sometimes almost all major colony-forms can be found together not enough known about factors controlling colony-forms in modern environments colony-forms represent alternative functional adaptive strategies that may be viable in many different environments representation of different colony forms has changed through the Phanerozoic
61 3. branch diameter as a depth indicator
62 Branch diameter variation within coral species as depth indicator shallow site intermediate site Coral Pocillopora damicornis (see Kaandorp 1999) deep site
63 Branching bryozoans (ramose, dendroid) Diaperoecia. Recent Pleistocene. Setana Fm Kuromatsunai, Japan
64 Heteropora pacifica Puget Sound, USA (Schopf et al. 1980)
65 three cyclostome species: Cinctipora elegans Diaperoecia purpurascens Erksonea sp. Taylor, Kuklinski & Gordon (2007)
66 Branch diameter does correlate with depth branch diameter (mm) r = -0.37, p <0.001 Error bars = 2 SD depth (metres)
67 Branch diameter does not correlate with depth branch diameter (mm) r = -0.13, p = Error bars = 2 SD depth (metres)
68 Branch diameter does not correlate with depth branch diameter (mm) r = -0.10, p = Error bars = 2 SD depth (metres)
69 4. zooid size as a temperature indicator
70 Cheilostome bryozoan zooid size is inversely correlated with ambient temperature at time of budding zooid length (microns) cheilostome bryozoan Electra pilosa C 12 C 18 C 22 C cultivation temperature
71 Cupuladria exfragminis upwelling (cold) zooids non-upwelling (warm) zooids Okamura et al. 2011
72 Cribrilina cryptooecium Cold Recent British Large zooids Warm Coralline Crag (Pliocene) Small zooids
73 MART analysis using within-colony variation in zooid size MART = mean annual range in temperature Aaron O Dea
74 Conopeum seurati Avonmouth, England
75 Conopeum seurati Avonmouth, England summer zooids
76 Conopeum seurati Avonmouth, England summer zooids winter zooids
77 seawater temperature zooid size summer winter summer
78 seawater temperature zooid size summer winter summer
79 seawater temperature zooid size summer winter summer
80 seawater temperature zooid size summer winter summer
81 seawater temperature zooid size summer winter summer
82 seawater temperature zooid size summer winter summer
83 seawater temperature zooid size summer winter summer
84 seawater temperature zooid size summer winter summer
85 seawater temperature zooid size summer winter summer
86 seawater temperature zooid size summer winter summer
87 seawater temperature zooid size summer winter summer
88 seawater temperature zooid size summer winter summer
89 zooid size seasonal climate high variance in zooid size
90 zooid size seasonal climate high variance in zooid size zooid size constant temperature no variance in zooid size
91 MART method
92 MART method randomly select 20 zooids
93 MART method randomly select 20 zooids avoid early zooids and areas of irregular growth
94 MART method randomly select 20 zooids avoid early zooids and areas of irregular growth measure zooid length and width
95 MART method randomly select 20 zooids avoid early zooids and areas of irregular growth measure zooid length and width calculate length x width (= area proxy)
96 MART method randomly select 20 zooids avoid early zooids and areas of irregular growth measure zooid length and width calculate length x width (= area proxy) calculate CV (= SD/mean x 100)
97 MART method randomly select 20 zooids avoid early zooids and areas of irregular growth measure zooid length and width calculate length x width (= area proxy) calculate CV (= SD/mean x 100) repeat in at least 4 more colonies
98 MART method randomly select 20 zooids avoid early zooids and areas of irregular growth measure zooid length and width calculate length x width (= area proxy) calculate CV (= SD/mean x 100) repeat in at least 4 more colonies
99 29 Recent cheilostome species MART = (b) where (b) is the mean intracolony CV of frontal area (length x width)
100 Contrasting seasonality between Caribbean and Pacific O Dea 2003
101 Contrasting seasonality between Caribbean and Pacific bryozoan estimate bryozoan estimate O Dea 2003
102 Early-Mid Pliocene North Atlantic Knowles et al. 2009
103 Early-Mid Pliocene North Atlantic Knowles et al. 2009
104 5. palaeolatitudinal distribution of bryozoan carbonates through time
105 Uniformitarianism Present is the Key to the Past' Charles Lyell ( )
106 Global distribution of carbonate deposition at the present-day
107 Global distribution of carbonate deposition at the present-day
108 Palaeolatitudinal Latitudinal distribution of distribution bryozoan-rich of carbonate bryozoan-rich sediments through sediments time Taylor & Allison Geology 26:
109 Palaeolatitudinal Latitudinal distribution of distribution bryozoan-rich of carbonate bryozoan-rich sediments through sediments time Tropics Taylor & Allison Geology 26:
110 Palaeolatitudinal Latitudinal distribution of distribution bryozoan-rich of carbonate bryozoan-rich sediments through sediments time Tropics Taylor & Allison Geology 26: post-palaeozoic are nearly all non-tropical
111 Palaeolatitudinal Latitudinal distribution of distribution bryozoan-rich of carbonate bryozoan-rich sediments through sediments time Tropics Taylor & Allison Geology 26: post-palaeozoic are nearly all non-tropical but Palaeozoic are pan-global
112 Palaeolatitudinal distribution of bryozoan-rich sediments post-palaeozoic Palaeozoic frequency palaeolatitude (degrees)
113 Palaeozoic post-palaeozoic Cincinnatian Series - Upper Ordovician, USA Tropical carbonate Coralline Crag - Pliocene, England Temperate carbonate
114 Bryozoan assemblage species richness vs latitude in the eastern (black dots) and western (white dots) North Atlantic no gradient of increasing species diversity into tropics (cf. many other phyla) bryozoans can be diverse in the tropics but have a relatively low biomass (as in Indonesian Cenozoic) Clarke & Lidgard Journal of Animal Ecology 69:
115 Postscript: analysis of annual growth bands as proxy for phytoplankton productivity in the Antarctic Cellarinella Recent Antarctica cf. tree rings
116 Summary and conclusions Akkeshi Bay
117 Summary and conclusions bryozoans at the present day inhabit many aquatic environments but are most diverse on continental shelves Akkeshi Bay
118 Summary and conclusions bryozoans at the present day inhabit many aquatic environments but are most diverse on continental shelves colony-form analysis must be treated cautiously Akkeshi Bay
119 Summary and conclusions bryozoans at the present day inhabit many aquatic environments but are most diverse on continental shelves colony-form analysis must be treated cautiously in some species relative branch diameter can be used as a palaeobathymetric indicator Akkeshi Bay
120 Summary and conclusions bryozoans at the present day inhabit many aquatic environments but are most diverse on continental shelves colony-form analysis must be treated cautiously in some species relative branch diameter can be used as a palaeobathymetric indicator zooid size correlates with temperature at time of budding and can be used to indicate relative palaeotemperature and also MART Akkeshi Bay
121 Summary and conclusions bryozoans at the present day inhabit many aquatic environments but are most diverse on continental shelves colony-form analysis must be treated cautiously in some species relative branch diameter can be used as a palaeobathymetric indicator zooid size correlates with temperature at time of budding and can be used to indicate relative palaeotemperature and also MART the latitudinal distribution of carbonate-producing bryozoans has changed through time Akkeshi Bay
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