Identifying AGRRA Corals: Part 4 Branching and Nodular Corals

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For images used in Part 4, our special thanks to: M. Brandt, K. Desai, I. Drysdale, D. Fenner, P. Humann, W. Precht, C. Rogers, R. Steneck, M. Vermeij, E. Weil, A. Yñiguez Reminder: What to Look for Underwater

Colony shape – massive (= mound, columnar, heavy plates), crust, plate, branching Colony size range – small to big Colony surface – bumpy, smooth, ridged Polyp size – small to big Polyp shape – round, elliptical, irregular, Y-shaped Polyp colour – brown, tan, yellow, olive, green, red Septal shape – fat, thin; smooth, toothed

Adapted from P.R. Kramer Reminder: AGRRA Coral Species The stony corals illustrated here are limited to species found in the wider Caribbean at depths (<20 m) typical of most AGRRA surveys. The names of some of these corals are changing as a result of modern research. More taxa are added as we gain underwater photographs of species that are rare and/or of geographically limited distributions. Expect periodic updates! Photographers who can enhance this collection are encouraged to contact Judy Lang at: [email protected] or [email protected]

For each species: (# in m and ft) = maximum colony size Remember: Coding Corals in AGRRA Surveys

Use the CARICOMP-based coral codes. The coral code for a genus (or occasionally a species complex) is the first 4 letters of its genus name and should be used whenever you are unsure of a coral’s species identity: MADR = Madracis The coral code for a species is the first letter of the genus name followed by the first 3 letters of its species name: MSEN = Madracis senaria

Codes are shown before names appear on the introductory slide for species that are commonly recorded in AGRRA surveys. Colony Boundaries in Branching Corals

Branching corals are easily broken and scattered, thus colony boundaries are often indistinct.

All species: tall thin polyps look fuzzy when expanded smooth branch surfaces are seen when polyps contract

© K. Marks Porites porites PPOR Porites porites PPOR thick (>2 cm/1 in), irregularly shaped branches, many PPOR with blunt tips elongate polyps often expand during the day light grey, cream, yellow- brown or blue

PPOR © R. Steneck

How differs from P. porites: branches are thinner, longer, more widely spaced, less irregular in shape, and with more rounded tips often darker colours (these are pale from bleaching)

© E. Weil P. porites P. furcata PPOR PFUR Porites divaricata PDIV thin (<1 cm/.4 in), short, widely-spaced branches, many subdivided near tip PDIV (“Y-shaped”) grey, yellow-brown to brown (clumps to ~ 30 cm/1 ft wide)

How differs from P. furcata: thinner, shorter, more widely- spaced branches, more often divided at tips fully grown colonies are smaller rare on fore reefs © C. Rogers Which is Which?

P. divaricata P. porites P. furcata PDIV PPOR PFUR Complications!

Some colonies look like “intermediates” of P. porites and P. furcata or of P. furcata and P. divaricata

If unsure of species identity, code as Porites “digitate” PDIG Madracis MADR

All species: small, round polyps are often exsert (“outies”)

polyps look fuzzy when tentacles are expanded

© K. Marks Madracis auretenra MAUR Madracis auretenra MAUR* thin, fragile, near-parallel, and densely packed MAUR* branches with blunt tips tentacles often expanded by day cream, yellow or yellow-brown

(usually to ~ 1.5 m/5 ft) © K. Marks

MAUR*

*MAUR, formerly called M. mirabilis, was described as a new species by Locke et al., 2007 (reference given on next-to-final slide). © K. Marks Madracis auretenra MAUR clumps many meters/10s of feet wide occur in sheltered habitats

How differs from Porites divaricata polyps are exsert branches are straighter and more closely set yellowish colours larger colonies when fully grown © K. Marks Which is Which?

P. divaricata M. auretenra PDIV MAUR Madracis decactis MDEC short, stubby nodules or lumpy crusts* MDEC MDEC small, exsert polyps, each usually with 10 © K. Marks conspicuous septa MDEC green, tan, grey, © K. Marks yellow-brown to dark brown (to ~ 1 m/3 ft)

*See Wells, 1973; encrusting colonies with 10 septa/polyp are also called Madracis decactis f. pharensis (Fenner, 1993, 1999) or morphospecies Madracis pharensis (Frade et al., 2010, and citations therein) (references given on next-to-final slide). Madracis decactis MDEC How knobby morph differs from M. auretenra + P. porites: forms stubby knobs, not branches darker colours fully grown colonies are smaller + from P. porites: septa are distinct (can be counted underwater)

M. decactis P. porites MDEC PPOR Which is Which?

© R. Steneck M. auretenra M. decactis MAUR MDEC Madracis senaria MSEN widely spaced polyps, each with about 6 large, exsert MSEN septa (and about 4 small septa) thick crusts that generally follow © M. Vermeij the contours of the substratum, can form conical-round columns MSEN green to orange-brown to dark brown (to ~ 30 cm/1 ft) MSEN

( to ~ 60 cm/2 ft) © M. Vermeij

MFOR MFOR

How differs from M. decactis polyps usually have 8 septa branching, not nodular, colonies that are larger when fully grown

polyps have 10 septa ovoid or rounded branches MCAR* often grow in the same plane and may have blunt tips brown with yellow-green polyp mouths © M. Vermeij (to ~ 2 m/6 ft)

MCAR* *M. carmabi was described as a new species by Vermeij et al., 2003, and is thought to be a hybrid of M. formosa and either M. decactis or M. pharensis sensu Frade et al., 2010 (references on

next-to-final slide). © M. Vermeij Madracis carmabi MCAR

How similar to M. formosa thick branches with blunt tips How similar to M. decactis + how differs from M. formosa: polyps have 10 conspicuous septa How differs from M. decactis: branching, not nodular, colonies that are larger when fully grown © M. Vermeij Which is Which?

Close-up Close-up

M. carmabi M. formosa MCAR MFOR Oculina diffusa ODIF short, somewhat twisted branches with ODIF large, distinct polyps yellow-brown, or may lack zooxanthellae (to ~ 30 cm/1 ft)

ODIF ODIF

How differs from M. auretenra: polyps are larger and protrude more above the skeleton

M. auretenra O. diffusa MAUR ODIF Acropora ACRO

All species: tiny (axial) polyps at the tips of actively growing branches are colourless (look white) © K. Marks Acropora palmata APAL lateral (radial) polyps are brown or yellow-brown (contain zooxanthellae)

APAL

ACER

APRO Acropora prolifera APRO

How the more “fused branch” forms differ from A. palmata: short branches don’t fuse into large thick masses fully grown colonies are smaller

APAL

APRO Acropora prolifera APRO

How the more “shrubby” and “spindly” forms differ from A. cervicornis: branches are shorter, straighter, more densely packed and may be thinner

How Millepora differs from the scleractinian stony corals: minute defensive polyps protrude from tiny pores in the colony surfaces

© K. Marks Millepora complanata Millepora complanata MCOM upright blades that rise above an encrusting base may have MCOM subdivided tips blades may interconnect to form complex “honeycombs” yellow, tan or yellow brown © K. Marks (to ~ 60 cm/2 ft tall) MCOM

MCOM

© K. Marks Millepora striata MSTR short, thin, narrow blades above an MSTR encrusting base have longitudinal MSTR “ribs” near their outer edges © K. Marks MSTR colonies vary greatly © D. Fenner in the extent to which the blades connect to form “paddles” or box-like structures

How differs from M. complanata: short, thin, narrow blades have “ribbed” tops fully grown colonies are smaller geographic distribution limited to southern and western Caribbean

© K. Marks Millepora squarrosa MSQU very short, honeycomb (box-like) structures with MSQU thick walls and rounded tops above an encrusting base © P. Humann tan to yellow-brown with reddish, pinkish or lavender tints MSQU geographic distribution limited to the northeastern and eastern Caribbean (to ~ 5 cm/2 in)

M. squarrosa M. complanata M. striata MSQU MCOM MSTR Millepora alcicornis MALC small, round to ovoid branches or thin crusts over other benthic animals (here MALC Gorgonia) and the substratum yellow to tan, branch tips white to ~ 45 cm/18 in

How differs from other species of Millepora small branches or thin crusts that grow over the substratum or other organisms

How differs from Oculina diffusa: minute polyps in tiny pores

M. complanata M. alcicornis MCOM MALC Which Is Which?

M. alcicornis O. diffusa MALC ODIF Complications! Some colonies look like “intermediates” of M. complanata and encrusting M. alcicornis (which is not surveyed in AGRRA coral transects but is included in AGRRA benthic point counts).

Recent research (e.g., Ruiz, 2009, and Tepper et al., 2012, references on final slide) has revealed the need for further studies to understand how the milleporids are related. References - Madracis Fenner, D.P. 1993. Species distinctions among several Caribbean stony corals. Bulletin of Marine Science 53: 1099-1116. Fenner, D. 1999. New observations on the stony coral (Scleractinia, Milleporidae, and Stylasteridae) species of Belize (Central America) and Cozumel (Mexico). Bulletin of Marine Science 64: 143-154. Frade, P.R., M.C. Reyes-Nivia, J. Faria, J.A. Kaandorp, P.C. Luttikhuizen, R.P.M. Bak. 2010. Semi-permeable species boundaries in the coral genus Madracis: Introgression in a brooding coral system. Molecular Phylogenetics and Evolution 57: 1072-1090. Locke, J.M., E. Weil, and K.A. Coates. 2007. A newly documented species of Madracis (Scleractinia: Pocilloporidae). Proceedings of the Biological Society of Washington 120: 214-226. Vermeij, M.J.A., O.E. Diekmann and R.P.M. Bak. 2003. A new species of scleractinian coral (Cnidaria, Anthozoa), Madracis carmabi n. sp. from the Caribbean. Bulletin of Marine Science 73: 679-864. Wells, J.W. 1973. Coral Reef Project–Papers in Memory of Dr. Thomas F. Goreau. 2. New and old scleractinian corals from Jamaica. Bulletin of Marine Science 23: 16-58. References - Millepora

Ruiz-Ramos, D.V. 2009. Morphological and genetic variation in the Caribbean species of the hydrocoral genus Millepora. M.Sc. Thesis. University of Puerto Rico. Abstract.

Tepper, C., L. Squiers, C. Hay, D. Gorgach, D. Friend, B. Black, B. Greenstein, K. Strychar. 2012. Cryptic species: A mismatch between genetics and morphology in Millepora. Marine Science 2: 57-65.