fleshy macroalgae share dominance with other organisms on ... · cnidarians (milleporans,...

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481 reefs at depths of 1-30 m (mean=10 ± 5.5 m) in the wider Caribbean were surveyed by the AGRRA protocols in 2011-2013. Any benthic group occupying at least 25% of the substratum at a site is here considered a spatial dominant. Sites can have more than one spatial dominant. All cover benthic point count values are “corrected” to remove any sand or mud as coral larvae are unlikely to settle here. W hen herbivores are scarce, fleshy macroalgae can expand over dead corals, outcompete live corals and prevent coral larvae from settling. But they are not the only players in the benthic spatial competition game. Fleshy Macroalgae Share Dominance with Other Organisms on Degraded Coral Reefs Judith C. Lang 1 , Philip A. Kramer 2 and Kenneth W. Marks 1 1 AGRRA (Atlantic and Gulf Rapid Reef Assessment), 2 TNC (The Nature Conservancy) 1. Detractors: Major Coral Competitors Fleshy macroalgae (FMA) Calcareous macroalgae (CMA) Turf Algal Sediment Mats (TAS) Peyssonnelids (PEY) Cyanobacteria (CYAN) “Aggressive” Invertebrates (AINV) FMA: highest mean cover (31±18.5%; max. = 84%), most abundant below 6 m and dominants at 60% of the sites. Often found growing with CMA. CMA: cover = 6.5±5% but a spatial dominant at only 3 sites. When added to FMA, dominance of macroalgae increased to 77% of sites. AINV: boring and certain overgrowing sponges, cnidarians ( milleporans, octocorals, zoanthids) and tunicates. Found in 82% of sites, collective mean cover of 2±2.5% (max. = 23%). PEY: overall uncommon (1.5±4%; max. = 56%), but rapidly expanding over dead Orbicella annularis on some reefs and dominants at 2 fore reefs. TAS: present in 77% of sites and most abundant in <6 m, averaging 6.5±5% cover, but spatial dominants at only 10% of the sites. CYAN: usually scarce (1.5±2.5% cover; max. = 16%). Thin films don’t harm corals which are killed by thick CYAN masses. TAS, close-up PEY CYAN boring sponge sponge close-up tunicate octocoral CMA FMA 5. Management Implications . Caribbean reefs need diverse herbivores–the echinoid Diadema antillarum (to consume TAS, PEY, MA) , large scarids (to help remove MA holdfasts), and acanthurids (to graze MA)–plus fewer nutrients, sediment and other pollutants on nearshore reefs (to facilitate coral health). blue parrotfish © S. Voegeli Diadema Thanks to: Ocean Research and Educational Foundation for support. Unless noted, all photos © K. Marks © C. Rogers blue tangs 2. Promotors: Major Reef Calcifiers Plus Crustose Coralline Algae (CCA) CCA: mean cover 10±8% (max. = 43%), spatial dominants in 6% of sites. Coral larval settlement sites; also spatial competitors of corals. CCA Sparse Turf Algae (TA) TA cover averaged 11.5±11% (max. = 56%), and declined with increasing depth. They were spatial dominants in 11% of sites. TA on CCA © R. Steneck SC Stony corals (SC) Live SC: averaged 14±10% (max. = 73%), were spatial dominants at 11% of sites. Highest cover was on bank reefs (26±18.5%) and at depths of <12 m. © K. Evans Grading Scale Poor Impaired Fair Good Benthic Index 1-1.5 2-2.5 3-3.5 4 3. Benthic Index: An attempt at integration For each site: Sum “reef promoters” cover: LC + CCA + sparse TA–a sign of herbivory and potential for coral larval settlement. Assign a score using the “promoter” threshold values below. Sum “reef detractors” cover: FMA + CMA + TAS + PEY + CYAN + AINV–all can displace LC and CCA. Assign a score using the “detractor” threshold values below. Average the two scores; assign a grade using the Benthic Index threshold values below. Note that BI, or any of the other indicators we assess on reefs, may reflect effects of both the ambient environment and human activities: some sites naturally are in better condition than others. Examples of Benthic Indices in the Wider Caribbean 4. Results. Impaired” is the most common BI score overall and at each depth range. The proportion of “fair” and “good” sites decreased with increasing depth. Most (n = 329) surveys were in fore reefs. Offshore banks (n = 29) had the highest proportion of “good” sites (~7%); intertidal crests (n=43) the highest proportion of “fair”(~30%). Sum Cover 1 (Poor) 2 (Impaired) 3 (Fair) 4 (Good) "Promoters" <15% 15-29.9% 30-59.9% 60% "Detractors" 60% 30-59.9% 15-29.9% <15% All Sites Stratified by Depth < 6 m Poor Impaired Fair Good n=137 77 28 28 4 6-<12 m Poor Impaired Fair Good n=174 110 16 45 3 18-30 m Poor Impaired 23 16 n=39 12-<18 m Poor Impaired Fair Good n=130 69 11 1 49 Poor Impaired Fair Good All Sites n=481 Impaired 280 Fair 55 8 Poor 128

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Page 1: Fleshy Macroalgae Share Dominance with Other Organisms on ... · cnidarians (milleporans, octocorals, zoanthids) and tunicates. Found in 82% of sites, collective mean cover of 2±2.5%

481 reefs at depths of 1-30 m (mean=10 ± 5.5 m) in the wider Caribbean were surveyed by the AGRRA protocols in 2011-2013. Any benthic group occupying at least 25% of the substratum at a site is here considered a spatial dominant. Sites can have more than one spatial dominant. All cover benthic point count values are “corrected” to remove any sand or mud as coral larvae are unlikely to settle here.

When herbivores are scarce, fleshy macroalgae can expand over dead corals, outcompete live corals and prevent coral larvae from settling. But they are not the only players in the benthic spatial competition game.

Fleshy Macroalgae Share Dominance with Other Organisms on Degraded Coral Reefs

Judith C. Lang1, Philip A. Kramer2 and Kenneth W. Marks11AGRRA (Atlantic and Gulf Rapid Reef Assessment), 2TNC (The Nature Conservancy)

1. Detractors: Major Coral CompetitorsFleshy macroalgae (FMA) Calcareous macroalgae

(CMA)Turf Algal Sediment Mats

(TAS)Peyssonnelids (PEY) Cyanobacteria (CYAN) “Aggressive” Invertebrates (AINV)

FMA: highest mean cover (31±18.5%; max. = 84%), most abundant below 6 m and dominants at 60% of the sites. Often found growing with CMA.

CMA: cover = 6.5±5% but a spatial dominant at only 3 sites. When added to FMA, dominance of macroalgae increased to 77% of sites.

AINV: boring and certain overgrowing sponges, cnidarians (milleporans, octocorals, zoanthids) and tunicates. Found in 82% of sites, collective mean cover of 2±2.5% (max. = 23%).

PEY: overall uncommon (1.5±4%; max. = 56%), but rapidly expanding over dead Orbicella annularis on some reefs and dominants at 2 fore reefs.

TAS: present in 77% of sites and most abundant in <6 m, averaging 6.5±5% cover, but spatial dominants at only 10% of the sites.

CYAN: usually scarce (1.5±2.5% cover; max. = 16%). Thin films don’t harm corals which are killed by thick CYAN masses.

TAS, close-up

PEY CYAN

boring sponge

sponge

close-up

tunicateoctocoral

CMA

FMA

5. Management Implications. Caribbean reefs need diverse herbivores–the echinoid Diadema antillarum (to consume TAS, PEY, MA), large scarids (to help remove MA holdfasts), and acanthurids (to graze MA)–plus fewer nutrients, sediment and other pollutants on nearshore reefs (to facilitate coral health).

blue parrotfish

© S. Voegeli

Diadema

Thanks to: Ocean Research and Educational Foundation for support. Unless noted, all photos © K. Marks

© C. Rogers

blue tangs

2. Promotors: Major Reef Calcifiers PlusCrustose Coralline Algae

(CCA)

CCA: mean cover 10±8% (max. = 43%), spatial dominants in 6% of sites. Coral larval settlement sites; also spatial competitors of corals.

CCA

Sparse Turf Algae (TA)

TA cover averaged 11.5±11% (max. = 56%), and declined with increasing depth. They were spatial dominants in 11% of sites.

TA on CCA

© R. Steneck

SC

Stony corals (SC)

Live SC: averaged 14±10% (max. = 73%), were spatial dominants at 11% of sites. Highest cover was on bank reefs (26±18.5%) and at depths of <12 m.

© K. Evans

Sum Cover 1 (Poor) 2 (Impaired) 3 (Fair) 4 (Good)"Promoters" <15% 15-29.9% 30-59.9% ≥60%"Detractors" ≥60% 30-59.9% 15-29.9% <15%

Grading Scale Poor Impaired Fair GoodBenthic Index 1-1.5 2-2.5 3-3.5 4

8

3. Benthic Index: An attempt at integrationFor each site: Sum “reef promoters” cover: LC + CCA + sparse TA–a sign of herbivory and potential for coral larval settlement. Assign a score using the “promoter” threshold values below.

Sum “reef detractors” cover: FMA + CMA + TAS + PEY + CYAN + AINV–all can displace LC and CCA. Assign a score using the “detractor” threshold values below.

Average the two scores; assign a grade using the Benthic Index threshold values below.

Note that BI, or any of the other indicators we assess on reefs, may reflect effects of both the ambient environment and human activities: some sites naturally are in better condition than others.

Examples of Benthic Indices in the Wider Caribbean

4. Results. “Impaired” is the most common BI score overall and at each depth range. The proportion of “fair” and “good” sites decreased with increasing depth. Most (n = 329) surveys were in fore reefs. Offshore banks (n = 29) had the highest proportion of “good” sites (~7%); intertidal crests (n=43) the highest proportion of “fair”(~30%).

Sum Cover 1 (Poor) 2 (Impaired) 3 (Fair) 4 (Good)"Promoters" <15% 15-29.9% 30-59.9% ≥60%"Detractors" ≥60% 30-59.9% 15-29.9% <15%

Grading Scale Poor Impaired Fair GoodBenthic Index 1-1.5 2-2.5 3-3.5 4

8

All Sites Stratified by DepthPoorImpairedFairGood

PoorImpairedFairGood

PoorImpairedFairGood

PoorImpaired

< 6 m

Poor Impaired Fair Good

6-<12 m

Poor Impaired Fair Good

12-<18 m

Poor Impaired Fair Good

18-30 m

Poor Impaired

n=137

77

28284

PoorImpairedFairGood

PoorImpairedFairGood

PoorImpairedFairGood

PoorImpaired

< 6 m

Poor Impaired Fair Good

6-<12 m

Poor Impaired Fair Good

12-<18 m

Poor Impaired Fair Good

18-30 m

Poor Impaired

n=174

110

1645

3

PoorImpairedFairGood

PoorImpairedFairGood

PoorImpairedFairGood

PoorImpaired

< 6 m

Poor Impaired Fair Good

6-<12 m

Poor Impaired Fair Good

12-<18 m

Poor Impaired Fair Good

18-30 m

Poor Impaired 23

16n=39

PoorImpairedFairGood

PoorImpairedFairGood

PoorImpairedFairGood

PoorImpaired

< 6 m

Poor Impaired Fair Good

6-<12 m

Poor Impaired Fair Good

12-<18 m

Poor Impaired Fair Good

18-30 m

Poor Impaired

n=13069

11 149

Poor 128 ALLImpaired 280Fair 55Good 8

All Sites

Poor Impaired Fair Good

All Sites

n=481

Impaired280

Fair 55

8Poor 128