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You are here: Home Volume VII September 2008 Feature Article: Coral Reproduction, Part Three: Stony Coral Sexuality, Reproduction Modes, Puberty Size, Sex Ratios and Life Spans

Feature Article: Coral Reproduction, Part Three: Stony Coral Sexuality, Reproduction Modes, Puberty Size, Sex Ratios and Life Spans

By Dana Riddle Posted Sep 14, 2008 08:00 PM Pomacanthus Publications, Inc.
Information on the reproductive habits of over 300 stony coral species (in almost 100 genera) is presented. It is the most complete single-source reference currently available.

The ultimate challenge of reefkeeping is the closing of our captive animals' life cycles. A fortunate few have witnessed a coral spawning in their aquariums and might be interested in what it would take to raise coral planula larvae. On the other hand, there may be dedicated hobbyists interested in captive spawning of a certain species. It is the goal of this series to assist those hobbyists. This time, we'll examine reproductive habits of stony corals. Information on the reproductive habits of over 300 stony coral species (in almost 100 genera) is presented. It is the most complete single-source reference currently available. This article will begin to offer information other factors (puberty age and size, life spans, etc.) and should begin to answer questions hobbyists might have.

image001.jpg

Figure 1. A female Sandalolitha robusta specimen spawns at the Waikiki Aquarium. Photo courtesy Dr. Bruce Carlson.

Witnessing coral reproduction, especially in an aquarium, is an exciting event. The first inclination is usually to find the camera and document the event. In the rush, some details may be (and often are) left unrecorded. Afterwards, many questions will inevitably arise. See the Figure 51 at the end of this article for suggestions on what to record.

There are several reasons why we don't hear of corals' sexual spawning in captivity, and the most likely reason is that few, if any, aquarists have specially tried to address issues pertinent to success. Another reason is that spawning events are simply not observed. It is also a fact that only a handful of dedicated aquarists have sufficiently researched requirements for captive breeding.

The goals of this third article on coral reproduction are simple. We'll examine the information available on stony coral sexuality and reproductive habits. This, in turn, will allow us to decide how many corals we'll likely need in order to have a pair (if necessary!). Sexual maturity is another issue. Unfortunately, we can not always judge the reproductive fitness of a coral simply by examining colony size - other factors are sometimes important (colony thickness is an important diagnostic tool in some small encrusting corals).

Be aware that benthic marine invertebrates as a group utilize almost every reproductive strategy imaginable, ranging from parthenogenesis (suggested by researchers to possibly occur in a few coral taxa) to other asexual means (including 'popping', 'dripping', 'budding', fragmentation, transverse fission, longitudal fission, stoloniferous growths, etc.) and, of course, sexual reproduction including broadcast spawning and brooding (both internally and externally). As usual, nature will continue to confound us as we realize that a particular coral species reproductive habit can, and often does, vary according to location and/or environmental conditions. However, this should not prevent us from reviewing reproduction data gathered from over 400 references (a complete reference list will appear at the end of this series).

To recap, our goals for this time are:

  1. Discuss tools to ID your stony coral to the species level.
  2. Understand its mode(s) of reproduction and with this information select the number and size of specimens to ensure a pair exists.
  3. Enable you to select an appropriate aquarium size. (Be aware that many corals' size at puberty might make them much too large for some home aquaria.)

Before continuing, perhaps a review is in order of terminology we'll use.

Glossary

Brooding or Brooder - Brooding
Where fertilized eggs are held internally (or sometimes on the surface of a parent colony) and are released as planula larvae. Brooder: A coral that uses the brooding reproductive process.
Fecundity
Fertility; ability to produce abundantly.
Gonochoric
Possessing distinct male and female colonies where offspring are a result of fusion of gametes. Also referred to as dioecious, or unisexual. Gonochorism occurs in ~25% of coral species examined (Richmond, 1997).
Hermaphroditic
Possessing both male and female reproductive organs, sometimes referred to as monoecious. Self-fertilization (also called 'selfing') is an uncommon hermaphroditic trait among corals.
Oocyte
An immature egg (ovum).
Parthenogenesis
Development of a new individual from an unfertilized egg. This results in a female clone and is thought to occur in many invertebrates (including soft corals, gorgonians and possibly stony corals) and some vertebrates.
Planula larvae
The free-swimming, ciliated stage of coral larvae.
Polyspermy
Where more than one sperm fertilizes an ovum.
Protandrous hermaphrodite
Where male sex organs mature before those of the female.
Protogynous
(proto=first; gynous = female) - Where female sex organs mature before those of the male.
Self-fertilization or 'Selfing'
Where gametes from a single parent colony are sufficient for successful reproduction. Relatively uncommon.

An Initial Recommendation

Unless you've witnessed a spawning of a particular coral species within your aquarium and want to investigate further, it is probably best to choose a coral that utilizes the brooding strategy. There are many advantages:

  1. Brooding coral species are generally smaller in size than their broadcasting counterparts.
  2. Planula larvae from brooders are relatively mature upon release and have the advantage of size.
  3. Some hermaphroditic brooders do not broadcast gametes. This lessens the chance of the aquarium's life support system suffering catastrophic failure.
  4. Brooded larvae sometimes contain zooxanthellae upon release from the parent colony, and the larvae do not have to obtain them from the surrounding environment.

Suspected Parthenogenesis

Parthenogenesis (definition above) is known to occur in a few invertebrates and even some vertebrates. It is suspected to occur in these stony corals:

  • Pocillopora damicornis (Permata et al., 2000)
  • Porites lobata (Fadllalah, 1983)
  • Porites lutea (Fadllalah, 1983)
  • Fungia scutaria (Krupp, 1983)

Self-fertilization ('selfing')

  • Acropora (Isopora) brueggemanni (Okubo et al., 2007)
  • Acropora tenuis (Heyward and Babcock, 1986)
  • Agaricia agaricites (Gleason et al., 2001).
  • Balanophyllia europaea (Goffredo et al., 2005).
  • Diploria strigosa (Hagman et al., 1998, in Alvarado et al., 2003)
  • Favia fragum (numerous references)
  • Goniastrea aspera (Heyward and Babcock, 1985)
  • Goniastrea favulus (Stoddart et al., 2004; Miller & Mundy, 2005)
  • Mycetophyllia species, suspected (Tirado, 2006)
  • Pocillopora damicornis (numerous references)
  • Porites astreoides (Gleason et al., 2001).
  • Seriatopora hystrix (Sherman, 2008)
  • Siderastrea radians (Neves et al., 2008)
  • Tubastraea coccinea (numerous references)

Gonochoric Brooding

Gonochoric brooding involves broadcast spawning by males and internal or external (surface) fertilization of oocytes, followed by internal or surface brooding:

  • Balanophyllia regia (Goffredo et al., 2005)
  • Dendrophyllia sp. (Babcock et al., 1986)
  • Goniopora queenslandiae (Yamazato et al., 1975, in Fadlallah, 1983)
  • Isophyllia dipsacea (Duerden, 1902, in Fadlallah, 1983)
  • Leptopsammia pruvoti (Kruzic et al., 2008)
  • Porites brighami (Richmond & Hunter, 1990)
  • Porites clavaria (Fadlallah, 1983)
  • Porites furcata (Soong, 1991)
  • Porites murrayensis (Neves, 1998)
  • Porites panamensis (Glynn et al., 2008)
  • Scolymia wellsi (suspected; Pitombo, 1992)
  • Tubastraea faulkerni (Babcock, 1986)

Brooders (stony corals)

  • Acrhelia horrescens (Kawaguti, 1941)
  • Acropora (Isopora) brueggemanni (see Richmond & Hunter, 1990)
  • Acropora corymbosa (Stimson, 1978)
  • Acropora (Isopora) cuneata (Wallace et al., 2007)
  • Acropora humilis (Stimson, 1978)
  • Acropora palawensis (see Richmond & Hunter, 1990)
  • Acropora (Isopora) palifera (Kojis, 1986)
  • Acropora striata (Stimson, 1978)
  • Acropora (Isopora) togianensis (Wallace et al., 2007)
  • Agaricia agaricites (Thornhill et al., 2006)
  • Alveopora daedalea (Schlesinger & Loya, 1985)
  • Balanophyllia europaea (Mezzomonaco et al., 2002)
  • Caryophyllia cyathus (Koch, 1897)
  • Cyphastrea ocellina (Stimson, 1978)
  • Dendrophyllia manni (see Richmond & Hunter, 1990)
  • Diploria strigosa (Hagman et al., 1998)
  • Euphyllia glabrescens (Kawaguti, 1941)
  • Favia fragum (Szmant, 1986)
  • Galaxea aspera (see Richmond & Hunter, 1990)
  • Goniopora queenslandiae (Loya, 1976)
  • Heliofungia actiniformis (see Richmond & Hunter, 1990)
  • Isophyllia dipsacea (Fadllalah, 1983)
  • Isopora togianensis (Wallace et al., 2007)
  • Madracis carmabi (Vermeij et al., 2004)
  • Madracis decactis (Vermeij et al., 2004)
  • Madracis formosa (Vermeij et al., 2004)
  • Madracis mirabilis (Vermeij et al., 2004)
  • Madracis pharensis (Vermeij et al., 2004)
  • Madracis senaria (Vermeij et al., 2004)
  • Manicina areolata (Johnson, 1992)
  • Mycetophyllia ferox (Babcock et al., 1986)
  • Pocillopora damicornis (Stimson, 1978)
  • Pocillopora elegans (Stimson, 1978)
  • Pocillopora verrucosa (Stimson, 1978)
  • Porites astreoides (Thornhill et al., 2006)
  • Porites panamensis (Glynn et al., 2008)
  • Scolymia wellsi (Pitombo, 1992)
  • Seriatopora caliendrum (Rinkevich & Loya, 1979)
  • Seriatopora hystrix (multiple references)
  • Siderastrea radians (Thornhill et al., 2006)
  • Tubastraea coccinea (Glynn et al., 2008)

Protandrous and Protogynous Hermaphrodites

Protandrous Hermaphrodites

  • Flabellum rubrum (Moseley, 1881, in Fadlallah, 1983)
  • Goniastrea favulus (Kojis and Quinn, 1981)
  • Stylophora pistillata (Loya, 1976)

Protogynous Hermaphrodite

  • Siderastrea radians (Duerden, 1902, in Fadlallah, 1983)

Degree of Difficulty

As we have seen, aquarists have been successful in propagating a number of coral species. There are a number of factors involved in these successes, beginning with the skill and dedication of the aquarist. Seasoned veterans often report 'easy' asexual reproduction of species difficult to maintain long-term in captivity (some Goniopora species are good examples, while there are no reports of reproduction of easily maintained species).

With that said (and avoiding the issues and degrees of difficult in husbandry of some coral species), I will unilaterally list my perception of how challenging captive breeding of corals could be (beginning with the least difficult):

  1. Those capable of Parthenogenesis.
  2. Programmed fragmentation. This method is essentially left to the coral (it is genetically programmed to produce colonies in this manner), and fragments 'drip' or 'pop' away. The only thing required from the hobbyist is securing the cloned colony to a suitable substrate.
  3. Intentional fragmentation. This technique requires little from the hobbyist. Small fragments, or nubbins, or snipped, clipped, sliced or cut away from brood stock and attached to any of a number of substrates for grow-out.
  4. Brooding coral colonies capable of self-fertilization or 'selfing'. Since 'selfing' is due to fertilization of an egg produced by a female by a sperm produced by a male within the same colony or sometimes the same polyp (hermaphroditic), these bisexual coral colonies can reproduce with only one colony present.
  5. Hermaphroditic brooding corals.
  6. Hermaphroditic broadcast spawners. In theory, only two coral colonies would be required to ensure cross-fertilization. However, many variables come into play such as sperm concentrations, polyspermy, egg buoyancy, impact of spawning on aquarium water quality, mode of zooxanthellae acquisition (vertical or horizontal), etc.
  7. Gonochoric broadcast spawners. In addition to the challenges presented by hermaphroditic broadcast spawners (above), we have to consider sex ratios (sometimes heavily skewed towards males) and the impact this has on sizing the aquarium.

Comments on Data Bases

While research this article, I found it best to maintain two data bases, and both are presented in this article. First, reproductive information about stony corals is listed in taxonomic order - by Family, Genus and Species along with comments, photographs, etc. After that, a quick reference sheet (along with the reference) presents reproductive information sans photos and comments (See Table 108).

A Note on Caribbean Corals

There have been rumors for a number of years that Caribbean corals would become legally available to hobbyists. After a number of false starts, this seems to be a reality. Rose corals (Manicina areolata) are available, and Siderastrea species may soon follow (Michael Janes, personnel communication. See www.aquatouch.com for details). Acropora palmata spats (grown from legally collected eggs and sperm from spawn slicks) are now in public aquaria. Perhaps this endangered species will someday be in home reef aquaria.

It has been estimated that 60% of Caribbean corals are brooders; some are of relatively small size and may be available in the future. For these reason, I have included Caribbean species in this article.

The Aquarium

Almost every article written on home-based fish propagation begins with aquarium selection. So, it is seems reasonable to start there, right? "Let's see - I've got that old 40 gallon breeder tank just gathering dust. I could use it…"

Whoa, cowboy! This line of reasoning has already dismissed the idea of logically thinking this project through from concept to reality. Fish breeders can choose the correct aquarium because they know much more about their fishes' reproductive habits than coral farmers know about their corals'. It is fairly easy to quickly research size of adult fish and how to sex them. So, let's start over.

Selecting Your Coral

Before you select your candidate coral, you should be able to identify it. There is a great deal of published research available (some of it presented in this series of articles), but utilization of this information depends upon correct identification of the coral. In some cases and with enough experience, this is simple. Some corals, such as the Elegance coral (Catalaphyllia jardineri), are not easily confused with any other species. Other corals will demonstrate a plasticity in shape and appearance according to environmental factors (water motion and lighting being two major influences), or naturally resemble very closely another coral species. For instance, there are several Acropora species that have pronounced corallites that appear to the casual viewer to be Acropora millepora, when in fact they are not.

Coral Identification Resources

Coral ID Software

Unless your stony coral identification skills are well-honed, I recommend the purchase of Veron's Coral ID software. See the Media Review on this excellent resource for stony coral species identification here (http://www.advancedaquarist.com/2008/4/review ).

Internet Resources

For Acropora species, there is an excellent free resource on the internet (www.coralsee.org).

Veron's Corals of the World is available for free at this website: http://whelk.aims.gov.au/coralsearch/coralid-search.php

Note this website is easier to use if you already know the coral ID to genus level. Then it is a matter of looking at each species. This can be a time consuming matter, and definitely is not as easy to reference as the print version.

If getting an ID to the genus level is acceptable, see CoralIdea (www.coralidea.com).

Books

There is no better book than Veron's Corals of the World (2000). See the review of this book here (http://www.advancedaquarist.com/2008/4/review).

With a little practice and a few basic laboratory tools (magnifying glass, dissecting microscope) you should be able to get a reasonable idea as to the correct species (it helps if you know where the corals were collected - keep this in mind if you order through any of the major distributors. They know!).

Can I Use Fragments? Effect of Damage (Fragmentation) on Fecundity

It is generally believed that growth, reproduction and maintenance are processes competing for a limited amount of energy. The effects of breakage on a particular colony depend upon the amount of damage, colony size, energy reserves (such as lipids), environmental conditions and other factors. Limited resources could be devoted to repair traumatized areas. It is also possible that corals will re-absorb oocytes/eggs to obtain their high-calorie yolks for use in tissue repair.

There are several published papers on the subject. Zakai (1997) found that breakage of Pocillopora damicornis exceeding 25% of colony area reduced planulation by 63% while having no effect on the size (but obviously the number) of planulae. Smith and Hughes, 1999 examined the effects of breakage on Acropora intermedia, Acropora millepora, Acropora hyacinthus and found that fecundity was greatly reduced.

However, fragmentation does not necessarily reduce fecundity. Daughter colonies produced by programmed fragmentation by Diaseris distorta are sexually functional at sizes as small as 1 cm2 (roughly ½" square). See Part 1 of this series for photos of Diaseris.

In general, fragments can be used, but expect fecundity to be reduced or delayed.

Polyp Anatomy

The sexual organs of corals are usually on mesenterial filaments within the polyp. See Figure 2.

image003.jpg

Figure 2. Anatomy of a 'typical' stony coral polyp. Eggs (oocytes) and spermaries (male gonads) - either separately or together - can be found on mesenteries within sexually mature polyps.

image005.jpg

Figure 3. Modes of coral reproduction. 'Selfing' and 'gonochoric brooding' are relatively uncommon, while the two forms of broadcast spawnings are not.

Sexing Your Coral

After selecting your target species, we can continue. Unfortunately, you can't just look at a coral and determine if it is male or female (unless you have the good fortune to witness it in the act of spawning, or have a small laboratory devoted to histology and can examine coral gonads). We'll need to understand a couple of terms before we continue.

Spawning Ratios - How Many of Each Species in an Aquarium?

Most corals are hermaphroditic and only in relatively few cases do we have to concern ourselves with the issue of sex ratios. Sex ratios, when available, are included in the initial reports (below). Be aware that sex ratios are not fixed and can vary from location to location, but this is the best information we currently have.

Notes on Colony Size, Sexual Maturity and Life Span

Colony size and sexual maturity has been linked in some cases, but many factors other than colony area should be considered. In some cases, it is not the size of the colony but the thickness of the coral's encrustation over the substrate that is the telling factor. In other cases, corals can asexually reproduce by fission, where a portion of the colony becomes isolated from the parent. In some cases, total mass of the colony (such as gorgonians) should be considered in lieu of colony height.

Generally, corals are a long-lived bunch and it is not uncommon to hear reports of corals believed to be hundreds of years of age. However, local patterns (number of storms, temperature, anthropogenic impacts, etc.) can limit survival rates.

On the other hand, brooding corals tend to have relatively brief lives. They mature at an earlier age and smaller size. They produce fewer young, but these tend to be well developed with a higher chance of survival. In some cases, brooders are opportunistic colonizers.

We'll now begin our review of stony coral reproductive habits. They are listed by in alphabetical order by Family, followed by Genus (also in alphabetical order). Information on species is presented in any of the Tables and some of the Figures.

For those wanting a quick reference, see the end of this article for Table 108 - The Quick and Easy Reference. There, corals are listed alphabetically by Genus.

Family Acroporidae

Genus Acropora (Staghorn and Antler Corals)

image007.jpg

Figure 4. Acropora species are easily propagated via intentional fragmentation by hobbyists, but there are reports of sexual spawning in aquaria. Photo by the author.

Table 1. Sexuality and reproductive Modes - Acropora species.
Taxon Sexuality Reproduction
Acropora aculeus Hermaphroditic Broadcast
Acropora acuminata Broadcast
Acropora anthoceris Hermaphroditic Broadcast
Acropora aspera Hermaphroditic Broadcast
Acropora austera Hermaphroditic Broadcast
Acropora caroliniana Broadcast
Acropora cerealis Hermaphroditic Broadcast
Acropora cervicornis Hermaphroditic Broadcast
Acropora clathrata Hermaphroditic Broadcast
Acropora corymbosa Hermaphroditic Brooder
Acropora cytheria Hermaphroditic Broadcast
Acropora danai Hermaphroditic Broadcast
Acropora delicatula Possibly sterile
Acropora delicatula See Acropora selago Broadcast
Acropora digitifera Hermaphroditic Broadcast
Acropora dilitata Hermaphroditic Broadcast
Acropora divaricata Hermaphroditic Broadcast
Acropora donei Hermaphroditic Broadcast
Acropora elseyi Broadcast
Acropora eurystoma Hermaphroditic Broadcast
Acropora exquisita Hermaphroditic Broadcast
Acropora florida Hermaphroditic Broadcast
Acropora formosa Hermaphroditic Broadcast
Acropora gemmifera Hermaphroditic Broadcast
Acropora glauca ? Broadcast
Acropora grandis Hermaphroditic Broadcast
Acropora grandulosa Hermaphroditic Broadcast
Acropora hemprichii Hermaphroditic Broadcast
Acropora horrida Hermaphroditic Broadcast
Acropora humilis Hermaphroditic Broadcast
Acropora humilis ? Brooder
Acropora hyacinthus Hermaphroditic Broadcast
Acropora hystrix Hermaphroditic Broadcast
Acropora intermeda Broadcast
Acropora irregularis Hermaphroditic Broadcast
Acropora jacquelineae Broadcast
Acropora kimbeensis
Acropora "kosurini-like" Broadcast
Acropora latistella Hermaphroditic Broadcast
Acropora lianae Broadcast
Acropora listeri Broadcast
Acropora longicyathus Hermaphroditic Broadcast
Acropora loripes Hermaphroditic Broadcast
Acropora lutkeni Hermaphroditic Broadcast
Acropora microclados Hermaphroditic Broadcast
Acropora microphthalma Hermaphroditic Broadcast
Acropora millepora Hermaphroditic Broadcast
Acropora monticulosa Hermaphroditic Broadcast
Acropora muricata Hermaphroditic Broadcast
Acropora nana Hermaphroditic Broadcast
Acropora nastua Hermaphroditic Broadcast
Acropora nobilis Hermaphroditic Broadcast
Acropora ocellata Hermaphroditic Broadcast
Acropora palawensis ? Brooder
Acropora palmata Hermaphroditic Broadcast
Acropora palmerae Broadcast
Acropora papillare Broadcast
Acropora plumosa Broadcast
Acropora polystoma Broadcast
Acropora pulchra Hermaphroditic Broadcast
Acropora robusta Hermaphroditic Broadcast
Acropora samoensis Hermaphroditic Broadcast
Acropora sarmentosa Hermaphroditic Broadcast
Acropora scandens Hermaphroditic Broadcast
Acropora secale Hermaphroditic Broadcast
Acropora selago Hermaphroditic Broadcast
Acropora smithi Hermaphroditic Broadcast
Acropora solitaryensis Hermaphroditic Broadcast
Acropora spathulata Broadcast
Acropora spicifera Broadcast
Acropora squarrosa Hermaphroditic Broadcast
Acropora striata ? Brooder
Acropora subulata Broadcast
Acropora surculosa Hermaphroditic Broadcast
Acropora tenuis Hermaphroditic Broadcast
Acropora tortuosa Hermaphroditic Broadcast
Acropora valenciennesi Hermaphroditic Broadcast
Acropora valida Hermaphroditic Broadcast
Acropora variabilis Hermaphroditic Broadcast
Acropora vaughani Broadcast
Acropora verweyi Hermaphroditic Broadcast
Acropora willisae Broadcast
Acropora yongei Hermaphroditic Broadcast

Puberty Age for Acropora Species

Wallace (1985) estimates Acropora granulosa, A. hyacinthus, A. loripes and A. valida first spawn when they are 4-5 years of age. See Table 2 for more information.

Table 2
Taxa Puberty Size Age Reference
Acropora spp. >20cm Guest et al., 2005
Acropora spp. 3 years Harrison & Wallace, 1990

Subgenus Isopora

Some controversy exists over 'subgenus' status of Isopora (some feel it should be elevated to genus status), but I'll present information as subgenus -as it is listed in the references - until this matter is resolved.

Table 3. Sexuality and reproductive Modes - Subgenus Isopora species.
Taxon Sexuality Reproduction
Acropora (Isopora) brueggemanni Hermaphroditic Brooder
Acropora (Isopora) cuneata Hermaphroditic Brooder
Acropora (Isopora) palifera Hermaphroditic Brooder
Acropora (Isopora) togianensis Hermaphroditic Brooder

Genus Anacropora

Table 4. Sexuality and reproductive Modes - Anacropora species.
Taxon Sexuality Reproduction
Anacropora matthai Broadcast

Genus Astreopora

Table 5. Sexuality and Reproductive Modes - Astreopora species.
Taxon Sexuality Reproduction
Astreopora gracilis Broadcast
Astreopora listeri Broadcast
Astreopora myriophthalma Hermaphroditic Broadcast
Astreopora randalli Hermaphroditic Broadcast

Genus Montipora

Montipora species are very popular among reef hobbyists and for good reason. As a group, they are hardy and can grow quickly. In addition, some species contain colorful fluorescent and chromoprotein pigments (see Figure 5).

image009.jpg

Figure 5. The fluorescence of Montipora danae. Photo by the author.

Table 6. Sexuality and Reproductive Modes - Montipora species. Note that M. dilitata, M. flabellate and M. patula are endemic to Hawaii, thus restricting their distribution within the pet trade.
Taxon Sexuality Reproduction
Montipora aequituberculata Hermaphroditic Broadcast
Montipora altasepta Hermaphroditic Broadcast
Montipora cactus Hermaphroditic Broadcast
Montipora capitata Hermaphroditic Broadcast
Montipora crassituberculata Hermaphroditic
Montipora digitata Hermaphroditic Broadcast
Montipora dilatata Hermaphroditic Broadcast
Montipora efflorescens Hermaphroditic Broadcast
Montipora effusa Broadcast
Montipora erythraea Hermaphroditic Broadcast
Montipora eydouxi Hermaphroditic Broadcast
Montipora faveolata Hermaphroditic Broadcast
Montipora flabellata Hermaphroditic Broadcast
Montipora floweri Hermaphroditic Broadcast
Montipora foliosa Hermaphroditic Broadcast
Montipora hispida Hermaphroditic Broadcast
Montipora informis Hermaphroditic Broadcast
Montipora monasteriata Hermaphroditic Broadcast
Montipora patula Hermaphroditic Broadcast
Montipora peltiformis Broadcast
Montipora ramosa Broadcast
Montipora samarensis Hermaphroditic
Montipora spumosa Hermaphroditic Broadcast
Montipora studeri Hermaphroditic Broadcast
Montipora tuberculosa Broadcast
Montipora turgescens Hermaphroditic Broadcast
Montipora turtlenesis Hermaphroditic Broadcast
Montipora venosa Hermaphroditic Broadcast
Montipora verrilli Hermaphroditic Broadcast
Montipora verrucosa Hermaphroditic Broadcast

Puberty Size of Montipora Specimens

Bassim (1997) reports that Montipora verrucosa (now M. capitata) colonies are sexually mature when they are 18 cm (7.2") on the longest axis.

Family Agariciidae

Members of Agariciidae are a mixed bag, reproductively speaking (Veron, 1986) - Agaricia species are brooders, while Pavona species are broadcast spawners. We don't have enough information on other genera to make generalizations. Unfortunately, there is presently a ban on collection of Agaricia species, but Pachyseris and Pavona specimens are common in reef aquaria.

Genus Agaricia(Lettuce Leaf Corals)

Table 7. Sexuality and Reproductive Modes - Agaricia species.
Taxon Sexuality Reproduction
Agaricia agaricites Hermaphroditic Brooder
Agaricia crassa ? Brooder
Agaricia fragilis ? Brooder
Agaricia grahame ? Brooder
Agaricia humilis Hermaphroditic Brooder
Agaricia lamarcki ? Brooder
Agaricia purpurea Hermaphroditic Brooder
Agaricia tenufolia ? Brooder
Agaricia undata ? Brooder

Puberty Size of Agaricia humilis

Agaricia species are found only in the Caribbean (Veron, 1986). As far as we know, all Agaricia species are hermaphroditic brooders. Agaricia humilis specimens are sexually mature when they are ~28mm in diameter (van Moorsel, 1983). A. humilis is known to planulate year round. See Figure 6 for details on adult colony sizes.

image012.jpg

Figure 6. Diameters of adult Agaricia species (Van Moorsel, 1981).

Genus Coeloseris

No information on reproductive habits available.

Genus Gardineroseris

Table 8. Sexuality of Gardineroseris planulata.
Taxon Sexuality Reproduction
Gardineroseris planulata Hermaphroditic

Genus Helioseris

Table 9. Reproductive Mode of Helioseris cucullata.
Taxon Sexuality Reproduction
Helioseris cucullata Brooder

Genus Leptoseris (Ridge Corals)

No information on reproductive habits available.

Genus Pachyseris


Table 10. Sexuality and Reproductive Modes of Pachyseris species.
Taxon Sexuality Reproduction
Pachyseris rugosa Gonochoric Broadcast
Pachyseris speciosa Gonochoric Broadcast

Genus Pavona (Pork Chop or Corrugated Corals)

image013.jpg

Figure 7. Pavona varians, from Hawaii. Photo by the author.

Table 11. Sexuality and Reproductive Modes of Pavona species.
Taxon Sexuality Reproduction
Pavona cactus Gonochoric ?
Pavona duerdeni Gonochoric Broadcast
Pavona explanata Gonochoric (?)
Pavona gigantea Gonochoric Broadcast
Pavona varians Gonochoric Broadcast

Family Astrocoeniidae

Genus Madracis

Madracis species are cosmopolitan - they are found in the Pacific (including Hawaii), Atlantic and Mediterranean.

Table 12. Sexuality and Reproductive Modes of Madracis species.
Taxon Sexuality Reproduction
Madracis carmabi Hermaphroditic Brooder
Madracis decactis Hermaphroditic Brooder
Madracis formosa Hermaphroditic Brooder
Madracis mirabilis Hermaphroditic Brooder
Madracis pharensis Hermaphroditic Brooder
Madracis senaria Hermaphroditic Brooder

Genus Palauastrea

Biologists seem to have overlooked this coral (it strongly resembles the common Porites cylindrica) and I have been unable to locate any information concerning its reproductive habits.

Genus Stephanocoenia

Table 13. Sexuality and Reproductive Modes of Stephanocoenia species.
Taxon Sexuality Reproduction
Stephanocoenia intercepta Gonochoric Broadcast
Stephanocoenia michelini Gonochoric Broadcast

Genus Stylocoeniella

Table 14. Sexuality of a Stylocoeniella species.
Taxon Sexuality Reproduction
Stylocoeniella sp. Gonochoric

Family Caryophylliidae

Genus Heterocyathus

A zooxanthellate coral (Veron, 2000). No information is available about reproductive habits.

Genus Caryophyllia

Table 15. Sexuality and Reproductive Modes of Caryophyllia species.
Taxon Sexuality Reproduction
Caryophyllia ambrosa (deep water) Hermaphroditic
Caryophyllia cornuformis (deep water) Hermaphroditic
Caryophyllia cyathus ? Brooder
Caryophyllia sequenzae (deep water) Hermaphroditic
Caryophyllia smithi Gonochoric Broadcast

Genus Goniocorella

Table 16. Sexuality of a Goniocorella species.
Taxon Sexuality Reproduction
Goniocorella dumosa (deep water) Gonochoric

Genus Lophelia

Table 17. Sexuality of a Lophelia species.
Taxon Sexuality Reproduction
Lophelia pertusa (deep water) Gonochoric ?

Family Dendrophylliidae

Some of the members of Dendrophylliidae are non-photosynthetic (they do not contain zooxanthellae). Since they are not autotrophic, some Balanophyllia, Dendrophyllia, Tubastraea and others require good water motion to deliver enough food, or require regular feedings. Those aquarists willing to meet the demands of these often brightly colored animals are often rewarded by their corals' captive spawnings (usually in the form of planula larvae).

Genus Balanophyllia

Balanophyllia europea (a Mediterranean coral) contains zooxanthellae. Other Balanophyllia species do not.

Table 18. Sexuality and Reproductive Modes of Balanophyllia species.
Taxon Sexuality Reproduction
Balanophyllia elegans Gonochoric Brooder
Balanophyllia europaea Hermaphroditic Brooder
Balanophyllia pruvoti Gonochoric
Balanophyllia regia Gonochoric (?) Brooder
Balanophyllia sp. Brooder

Age of Puberty, Sex Ratio and Life Span for Balanophyllia species

Balanophyllia elegans colonies are capable of reproduction (brooding) at 1.5 years of age and have an expected life span of about 10 years (Gerrodette, 1981).

The gonochoric Balanophyllia pruvoti has a sex ratio of 1:1 (Radetic et al., 2002).

Genus Cladopsammia

Table 19. Sexuality and Reproductive Modes of Cladopsammia rolandi.
Taxon Sexuality Reproduction
Cladopsammia rolandi Hermaphroditic Brooder

Genus Dendrophyllia

Table 20. Sexuality and Reproductive Modes of a Dendrophyllia species.
Taxon Sexuality Reproduction
Dendrophyllia nigrescens See Tubastraea spp.
Dendrophyllia sp. Gonochoric Brooder

Genus Duncanopsammia

No information available on reproduction of the single member of this genus - Duncanopsammia axifuga (Veron, 1986).

Genus Enallopsammia

Table 21. Sexuality and Reproductive Mode of Enallopsammia rostrata.
Taxon Sexuality Reproduction
Enallopsammia rostrata( deep water) Gonochoric

Genus Leptopsammia (Sunset Cup Coral)

Table 22. Sexuality and Reproductive Mode of Leptopsammia pruvoti.
Taxon Sexuality Reproduction
Leptopsammia pruvoti Gonochoric Brooder

Genus Heteropsammia

These small (25mm diameter) corals live at depths of ~25 meters or more and are free-living on soft bottoms (Veron, 2000). They may or may not contain zooxanthellae.

Table 23. Sexuality and Reproductive Modes of Heteropsammia species.
Taxon Sexuality Reproduction
Heteropsammia aequicostatus Gonochoric Broadcast
Heteropsammia cochlea Gonochoric Broadcast

Genus Rhizopsammia (Wellington's Solitary Coral)

Table 24. Sexuality and Reproductive Modes of Rhizopsammia minuta.
Taxon Sexuality Reproduction
Rhizopsammia minuta Brooder

Genus Tubastraea(Sun Corals)

image015.jpg

Figure 9. This Tubastraea colony originated from a planula larva that settled in an area of high flow - an overflow in a reef aquarium.

Tubastraea (or perhaps Dendrophyllia) specimens reproduce readily in an aquarium when conditions are right.

Fully developed planula larvae are released and the number of reports of this occurrence in aquaria rivals that of another brooding coral - Pocillopora damicornis.

Table 25. Sexuality and Reproductive Modes of Tubastraea species.
Taxon Sexuality Reproduction
Tubastraea aurea See Tubastraea coccinea
Tubastraea coccinea Hermaphroditic Brooder
Tubastraea faulkneri Gonochoric Brooder

Puberty Size and Age of Tubastraea coccinea

Tubastraea coccinea (Glynn et al., 2008) Reproductive at 1.5 years, equating to colony size of ~5 cm in diameter, although Glynn et al., 2008 reports colonies can be reproductive when only 2 to 10 polyps are present (possibly due to stoloniferous growths). Paz-García et al. (2007) also reports reproduction in colonies only 5 cm in diameter (see Figure 11).

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Figure 10. These Tubastraea and Balanophyllia species are sexually mature at an early age.

Genus Turbinaria (Pagoda Corals)

Veron, 1986 states that Turbinaria species are all gonochoric broadcast spawners.

image019.jpg

Figure 11. The distinctive yellow-green coloration of Turbinaria reniformis is visible in this specimen. Photo by the author.

Table 26. Sexuality and Reproductive Modes of Turbinaria species.
Taxon Sexuality Reproduction
Turbinaria frondens Gonochoric
Turbinaria mesenterina Broadcast
Turbinaria reniformis Gonochoric Broadcast
Turbinaria sp. Gonochoric Broadcast

Family Euphyllidae

Family Euphylliidae contains some of the most popular aquarium corals including Euphyllia species (commonly called Anchor, Hammer, and Frogspawn corals) as well the Elegance coral (Catalaphyllia jardinei), Bubble and Grape corals (genera Plerogyra and Physogyra, respectively) and the Fox coral (Nemenzophyllia).

Euphylliidae corals, as a group, use various reproductive strategies as the following tables demonstrate.

Genus Catalaphyllia (Elegance Coral)

There is only one known Catalaphyllia species.

Table 27. Sexuality and Reproductive Modes of Catalaphyllia jardeni.
Taxon Sexuality Reproduction
Catalaphyllia jardinei Gonochoric Broadcast

Genus Euphyllia (Hammer, Anchor, and Frogspawn Corals)

image021.jpg

Figure 12. Euphyllia ancora, a gonochoric broadcast spawner. Photo by the author.

Euphyllia species are of separate sexes (gonochoric) and broadcast gametes, except for the hermaphroditic brooder E. glabrescens. There are reports of E. glabrescens planulating and successful settlements of larvae within aquaria (Mitch Carl, personal communication).

Table 28. Sexuality and Reproductive Modes of Euphyllia Species.
Taxon Sexuality Reproduction
Euphyllia ancora Gonochoric Broadcast
Euphyllia divisa Gonochoric Broadcast
Euphyllia glabrescens Hermaphroditic Brooder
Euphyllia parancora Gonochoric? Broadcast
Euphyllia rugosa Brooder
image023.jpg

Figure 13. Release of planula larvae by a Euphyllia glabrescens at the Henry Doorly Zoo, Omaha, Nebraska. Photo courtesy Mitch Carl.

Maximum Colony Sizes of Brooding and Broadcasting Euphyllia Species

image026.jpg

Figure 14. Euphyllia glabrescens is a hermaphroditic brooder, and is smaller than its broadcasting relatives. Note that 100 cm diameter may not be the largest colony size, it is the largest category listed by the researcher (Soong, 1993).

Genus Nemenzophyllia (The Fox Coral)

Some of the corals sold as Nemenzophyllia are likely Plerogyra discus. Nemenzophyllia is often listed as rare, and no information is available on its reproductive habits.

image027.jpg

Figure 15. Nemenzophyllia turbida or Plerogyra discus? Only its taxonomist knows for sure. Photo by the author.

Genus Physogyra (Grape Coral)

The Grape coral is commonly seen in reef aquaria. It is a gonochoric, broadcast spawner.

Table 29. Sexuality and Reproductive Mode of Physogyra lichtensteini.
Taxon Sexuality Reproduction
Physogyra lichtensteini Gonochoric Broadcast

GenusPlerogyra (Bubble Coral)

The Bubble coral is another popular reef aquarium inhabitant. It is probably gonochoric, and definitely a broadcast spawner.

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Figure 16. Hobbyists have reported that Bubble Corals (Plerogyra sinuosa) are broadcast spawners. Photo courtesy of Steve Ruddy.

Table 30. Reproductive Mode of Plerogyra sinuosa.
Taxon Sexuality Reproduction
Plerogyra sinuosa Broadcast

Genus Solenosmilia

Table 31. Sexuality and Reproductive Modes of Solensomilia variabilis.
Taxon Sexuality Reproduction
Solensomilia variabilis Gonochoric Broadcast (?)

Family Faviidae

Genus Astreosmilia

Limited to the western Indian Ocean (Veron, 1986), few researchers have paid much attention to these corals, and no information on spawning habits is available.

Genus Australogyra

Table 32. Sexuality and Reproductive Mode of Australogyra zelli.
Taxon Sexuality Reproduction
Australogyra zelli Hermaphroditic Broadcast

Genus Barabattoia

Table 33. Sexuality and Reproductive Mode of Barabattoia amicorum.
Taxon Sexuality Reproduction
Barabattoia amicorum Hermaphroditic Broadcast

Genus Caulastrea (Candy Cane Corals)

Table 34. Sexuality and Reproductive Mode of Caulastrea furcata.
Taxon Sexuality Reproduction
Caulastrea furcata Hermaphroditic Broadcast

Genus Cladocora (Tube Corals)

Cladocora specimens are often found on Florida live rock but their patchy spots or clusters of small polyps are not particularly spectacular. Some do not possess zooxanthellae making their small tentacles colorless. Their 'tubes' are ~1/8" in diameter.

Table 35. Sexuality and Reproductive Mode of Cladocora caespitosa.
Taxon Sexuality Reproduction
Cladocora caespitosa Hermaphroditic Broadcast

Genus Colpophyllia(Grooved Brain Coral)

This genus is found only in the Atlantic (Veron, 1986).

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Figure 17. Colpophyllia species. Photo courtesy Jake Adams and www.coralidea.com

Table 36. Sexuality and Reproductive Modes of Colpophyllia species.
Taxon Sexuality Reproduction
Colpophyllia amaranthus Broadcast
Colpophyllia breviseralis Broadcast
Colpophyllia natans Hermaphroditic Broadcast

Genus Cyphastrea

Table 37. Sexuality and Reproductive Modes of Cyphastrea species.
Taxon Sexuality Reproduction
Cyphastrea chalcidicum Hermaphroditic Broadcast
Cyphastrea microphthalma Hermaphroditic Broadcast
Cyphastrea ocellina Hermaphroditic Brooder
Cyphastrea seraila Hermaphroditic Broadcast

Maximum Colony Sizes of Various Cyphastrea Species

Information on maximum colony sizes of various Cyphastrea confirms once again that brooding corals are smaller in size than their broadcasting relatives. See Figure 18.

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Figure 18. The brooder Cyphastrea ocellina has a smaller adult colony size than the Cyphastrea species that are broadcast spawners.

Genus Diploastrea

Table 38. Sexuality and Reproductive Mode of Diploastrea heliopora.
Taxon Sexuality Reproduction
Diploastrea heliopora Gonochoric Broadcast

Genus Diploria (Brain Corals)

This genus is found only in the Atlantic (Veron, 1986), and it is possible that specimens could hitchhike in on legally collected live rock.

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Figure 19. Adventurer and reef hobbyist John Dawe watches a Diploria spawn at the Flower Garden Banks, Gulf of Mexico. Egg/sperm bundles are visible as white dots throughout the picture. Photo courtesy of Michael P. Janes.

Table 39. Sexuality and Reproductive Modes of Diploria species.
Taxon Sexuality Reproduction
Diploria clivosa Hermaphroditic Broadcast
Diploria labyrinthiformis Hermaphroditic Broadcast
Diploria labyrinthiformis Hermaphroditic Brooder
Diploria strigosa Hermaphroditic Broadcast
Diploria strigosa Hermaphroditic Brooder

Sizes and Sexual Maturity of Diploria clivosa and D. strigosa

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Figure 20. Diploria clivosa is a hermaphroditic broadcast spawner.

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Figure 21. Diploria strigosa is a hermaphroditic brooder.

Table 40. Diploria Puberty Sizes and Ages
Taxa Puberty Size Age Reference
Diploria clivosa >100 cm² 7-9 years Soong, 1992
Diploria strigosa >100 cm² 7-9 years Soong, 1992

Genus Echinopora

Table 41. Sexuality and Reproductive Modes of Echinopora species.
Taxon Sexuality Reproduction
Echinopora gemmacea Hermaphroditic Broadcast
Echinopora horrida Hermaphroditic Broadcast
Echinopora lamellosa Hermaphroditic Broadcast
Echinopora pacificus Hermaphroditic Broadcast

Genus Erythrastrea

This genus' distribution is limited to the Red Sea (Veron, 1986) and no information on its reproductive habits is available.

Genus Favia (Star Corals)

Favia corals have been a mainstay in the reef aquarium hobby for many years. There are scattered reports of reproduction in aquaria.

image041.jpg

Figure 22. A Favia specimen releases an egg or sperm bundle in the aquarium of master aquarist Tony Vargas. Photo courtesy of Tony Vargas. This Favia has been maintained in captivity since 2000, and is 8 inches in diameter. Spawning occurred in June 7, 2004 in the early morning, and the event lasted for ~1 hour. This coral had recently endured a move from New York to Florida.

Table 42. Sexuality and Reproductive Modes of Favia species.
Taxon Sexuality Reproduction
Favia bennettae Hermaphroditic Broadcast
Favia doreyensis Hermaphroditic Broadcast
Favia favus Hermaphroditic Broadcast
Favia favus Gonochoric Broadcast
Favia favus Hermaphroditic Broadcast
Favia fragum Hermaphroditic Brooder
Favia helianthoides Hermaphroditic Broadcast
Favia laxa Broadcast
Favia lizardensis Hermaphroditic Broadcast
Favia matthai Hermaphroditic Broadcast
Favia pallida Hermaphroditic Broadcast
Favia rotumana Broadcast
Favia speciosa Hermaphroditic Broadcast
Favia stelligera Hermaphroditic Broadcast
Favia veroni Hermaphroditic Broadcast
Table 43. Size and maturity of several Favia species.
Taxa Puberty Size Age Reference
Favia doryensis ? 8 years Connell, 1973
Favia favus ~3.5 cm² 4 years Oren et al., 2001
Favia fragum 2- 4 cm² 1.5 - 2 years Soong, 1992

Notes on Favia Puberty Size and Age

  • Favia doryensis specimens are sexually mature at 8 years of age (Connell, 1974).
  • Favia fragum has reproduced in aquaria. It is a small coral (no more than 2" in diameter and broods its young).
image044.jpg

Figure 23. Favia fragum is known to reproduce in public aquaria. It is a brooder, matures at a small size (~ 50mm in diameter) and can be very attractive. Unfortunately, it is an Atlantic coral and is not readily available to hobbyists.

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Figure 24. The trend continues… the brooding coral F. fragum is smaller than its broadcast spawning counterparts. Note that 100 cm diameter may not be the largest colony size, it is the largest category listed by the researcher (Soong, 1993).

Genus Favites

Table 44. Sexuality and reproduction modes of Favites species.
Taxon Sexuality Reproduction
Favites abdita Hermaphroditic Broadcast
Favites bennettae Hermaphroditic Broadcast
Favites chinensis Hermaphroditic Broadcast
Favites complanata Hermaphroditic Broadcast
Favites flexuosa Hermaphroditic Broadcast
Favites halicora Hermaphroditic Broadcast
Favites pentagona Hermaphroditic Broadcast
Favites russelli Hermaphroditic Broadcast

Genus Goniastrea (Honeycomb Coral)

Table 45. Sexuality and reproduction modes of Goniastrea species.
Taxon Sexuality Reproduction
Goniastrea aspera Hermaphroditic Broadcast
Goniastrea aspera Hermaphroditic Brooder
Goniastrea australensis Hermaphroditic Broadcast
Goniastrea edwardsi Hermaphroditic Broadcast
Goniastrea favulus Hermaphroditic Broadcast
Goniastrea favulus Protandrous Hermaphrodite
Goniastrea palauensis Hermaphroditic Broadcast
Goniastrea pectinata Hermaphroditic Broadcast
Goniastrea retiformis Hermaphroditic Broadcast

Puberty Size and Age of Goniastrea Specimens

  • Goniastrea aspera - less than 3"in diameter have only a slim chance of containing gonads, and proportion of mature colonies increases with colony size (Babcock, 1984)
  • Goniastrea aspera and G. favulus reached reproductive age in about 5 years (Babcock, 1991).

Genus Leptastrea

Table 46. Sexuality and reproduction modes of Leptastrea species.
Taxon Sexuality Reproduction
Leptastrea bottae Gonochoric Broadcast
Leptastrea purpurea Gonochoric Broadcast

Genus Leptoria

Table 47. Sexuality and reproduction mode of Leptoria phyria.
Taxon Sexuality Reproduction
Leptoria phryia Hermaphroditic Broadcast

Genus Manicina(Common Atlantic Rose Coral)

These beautiful Caribbean corals are being legally reared and are beginning to find there way to market in the U.S. Their small size and reproductive habits make them an idea candidate for captive propagation efforts. See this website for details on obtaining Manicina specimens: www.aquatouch.com

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Figure 25. The Caribbean Rose Coral (Manicina areolata) is now legally available for purchase by hobbyists. Photo courtesy Jake Adams and www.coralidea.com

Table 48. Sexuality and reproduction mode of Manicina areolata.
Taxon Sexuality Reproduction
Manicina areolata Hermaphroditic Brooder

Comment on M. areolata Adult Size

Adult Manicina specimens are ~3 inches long.

Genus Montastrea (Boulder Coral)

Montastrea corals are found in the Atlantic and Pacific Oceans.

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Figure 26. Montastrea cavernosa. Photo courtesy Jake Adams and www.coralidea.com

Table 49. Sexuality and reproduction modes of Montastrea species.
Taxon Sexuality Reproduction
Montastrea magnistellata Hermaphroditic Broadcast
Montastrea annularis Hermaphroditic Broadcast
Montastrea annuligera Hermaphroditic Broadcast
Montastrea cavernosa Gonochoric Broadcast
Montastrea curta Hermaphroditic Broadcast
Montastrea faveolata Hermaphroditic Broadcast
Montastrea franksi Hermaphroditic Broadcast
Montastrea valenciennesi Hermaphroditic Broadcast
image052.jpg

Figure 27. The Caribbean faviid Montastrea cavernosa has a sharply defined sexual maturity size.

Age of Puberty for Montastrea annularis

Szmant (1986) estimates the age of puberty for Montastrea annularis is 5 to 6 years.

Genus Moseleya

Table 50. Sexuality and reproduction modes of Moseleya latistelata.
Taxon Sexuality Reproduction
Moseleya latistellata Hermaphroditic Brooder

Genus Oulastrea (Zebra Coral)

Table 51. Sexuality and reproduction modes of Oulastrea crispa.
Taxon Sexuality Reproduction
Oulastrea crispa Hermaphroditic Broadcast
Oulastrea crispa Hermaphroditic Brooder

Puberty Size of Oulastrea crispa

Colonies greater than 20mm in diameter are sexually mature (Lam, 2000).

Genus Oulophyllia

Table 52. Sexuality and reproduction modes of Oulophyllia species.
Taxon Sexuality Reproduction
Oulophyllia bennettae Hermaphroditic Broadcast
Oulophyllia crispa Hermaphroditic Broadcast

Genus Parasimplastrea

There is only one species (Parasimplastrea sheppardi). It has a very limited distribution, and no information is available on its reproductive habits.

Genus Platygyra

Table 53. Sexuality and reproduction modes of Platygyra species.
Taxon Sexuality Reproduction
Platygyra contorta Hermaphroditic Broadcast
Platygyra daedalea Hermaphroditic Broadcast
Platygyra lamellina Hermaphroditic Broadcast
Platygyra pini Hermaphroditic Broadcast
Platygyra ryukuensis Hermaphroditic Broadcast
Platygyra sinensis Hermaphroditic Broadcast
Platygyra verweyi Hermaphroditic Broadcast

Puberty Age of Platygyra sinensis

Platygyra sinensis reaches a reproductive state at ~5 years of age (Babcock, 1991).

Genus Plesiastrea

Table 54. Sexuality and reproduction modes of Plesiastrea verispora.
Taxon Sexuality Reproduction
Plesiastrea verispora Broadcast

Genus Solenastrea

Solenastrea is an Atlantic species (Veron, 1986).

Table 55. Reproduction modes of Solenastrea species.
Taxon Sexuality Reproduction
Solenastrea boumoni Broadcast
Solenastrea hyades Broadcast

Family Flabelliidae

Genus Flabellum

Table 56. Sexuality of Flabellum species.
Taxon Sexuality Reproduction
Flabellum alabastrum (deep water) Gonochoric
Flabellum angulare (deep water) Gonochoric
Flabellum curvatum (deep water) Gonochoric
Flabellum impensum (deep water) Gonochoric
Flabellum japonicum Gonochoric
Flabellum rubrum Protandrous Hermaphrodite
Flabellum rubrum Hermaphroditic
Flabellum thouarsii (deep water) Gonochoric

Notes on Flabellum rubrum Sexuality and Size

Flabellum rubrum - small colonies (13-20mm) are males; larger colonies are hermaphroditic (Gardiner, 1902, in Fadlallah, 1983).

Genus Gardineria

Azooxanthellate. No information on reproduction is available.

Genus Monomyces

No information on reproduction is available.

Genus Placotrochus

No information on reproduction is available.

Family Fungiidae

image053.jpg

Figure 28. A female Fungia scutaria spawning at the Hawaii Institute of Marine Biology, Oahu, Hawaii. Note the tiny orange eggs being released. Photo courtesy of Jake Adams.

As a rule, all Fungiids seen to be gonochoric with the exception of one report of Heliofungia actiniformis being a brooder.

Genus Cantharellus

This Fungiid has a wide geographical range, but is rarely seen. It sometimes lives attached to the substrate (Veron, 2000). No information is available on reproductive habits.

Genus Ctenactis

No information is available on reproductive habits.

Genus Cycloseris

YouTube has some interesting video clips showing a Cycloseris (tentative ID) broadcast spawning. It is tempting to state that Cycloseris, like other Fungiids, is a gonochoric broadcast spawner.

Genus Danafungia

No information is available on reproductive habits.

Genus Diaseris

Table 57. Sexuality and reproduction mode of Diaseris distorta.
Taxon Sexuality Reproduction
Diaseris distorta Gonochoric Broadcast
image056.jpg

Figure 29. Sex ratios of Diaseris distorta in Galapagos Islands, Ecuador, where there are 5 males to every female colony.

Puberty Size and Sex Ratios of Diaseris distorta

Diaseris distorta (Colley et al., 2000) - Satellite colonies possess sex organs when colony size is ~1 cm2, while colonies produced sexually do not reach puberty until they are of larger size . Male colonies outnumber females by a ratio of 5 to 1 (in Ecuador).

Genus Fungia (Mushroom Coral)

Table 58. Sexuality and Reproduction Modes of Fungia species.
Taxon Sexuality Reproduction
Fungia actiniformis See Heliofungia actiniformis
Fungia concinna Gonochoric Broadcast
Fungia fungites Gonochoric Broadcast
Fungia granulosa Gonochoric Broadcast
Fungia horrida Broadcast
Fungia paumotensis Gonochoric Broadcast
Fungia repanda Gonochoric Broadcast
Fungia scutaria Gonochoric Broadcast
Fungia scutaria Parthenogenic?
Fungia sp. (Okinawa) Gonochoric Broadcast
image058.jpg

Figure 30. All Fungia species are gonochoric broadcast spawners. Sex ratios vary among species, and then even with environmental conditions. Based on information from Kramarsky-Winter and Loya (1998) and others.

Puberty Size, Age and Lifespan of Fungia Species

Fungia concinna and Fungia fungites are capable of reproduction at 4 years of age (equaling a size of ~6 cm in diameter; Harrison and Wallace, 1990). This information is in good agreement with that of Goffredo and Chadwick-Furman (2004) who report Fungia scutaria specimens are approximately 5 years of age when obtaining a diameter of 7 cm.

Further, these researchers report F. scutaria specimens are about 14 years old when they obtain a diameter of 22 cm. The estimated life span of F. scutaria is about 50 years.

Table 59. Size at puberty of various Fungiids.
Taxa Puberty Size Age Reference
Fungia coccinea 6 cm diameter 4 years Harrison & Wallace, 1990
Fungia fungites 6 cm diameter 4 years Harrison & Wallace, 1990
Fungia grandulosa 5.5 cm diameter ? Kramarsky-Winter & Loya, 1998
Fungia scutaria (male) 2.5 - 6 cm diameter ? Kramarsky-Winter & Loya, 1998
Fungia scutaria (female) >9 cm diameter ? Kramarsky-Winter & Loya, 1998
Heliofungia actiniformis 8 cm diameter 10 years Connell, 1973
image060.jpg

Figure 31. Fungiids can be reproductive over a broad range of sizes.

Genus Fungicyathus

This azooxanthellate coral can live at extreme depths - over 6,000 meters - the deepest of any coral (Veron, 2000). Needles to say, this coral is not likely to be found in your average reef aquarium!

Table 60. Sexuality of Fungicyathus species.
Taxon Sexuality Reproduction
Fungicyathus crispa Gonochoric
Fungicyathus fragilis Gonochoric
Fungicyathus marenzelleri Gonochoric

Genus Halomitra

No information available.

Genus Heliofungia (Plate Coral)

Table 61. Sexuality and Reproduction Modes of Heliofungia actiniformis.
Taxon Sexuality Reproduction
Heliofungia actiniformis Hermaphroditic (?) Brooder
Heliofungia actiniformis Gonochoric Broadcast

Heliofungia Puberty Size and Age

Heliofungia actiniformis is reproductive at 10 years of age with a corresponding diameter of 8cm (Connell, 1973).

Genus Herpetoglossa

Table 62. Sexuality and Reproduction Mode of Herpetoglossa simplex.
Taxon Sexuality Reproduction
Herpetoglossa simplex Gonochoric Broadcast

Genus Herpolitha

Table 63. Sexuality and Reproduction Mode of Herpolitha limax.
Taxon Sexuality Reproduction
Herpolitha limax Gonochoric Broadcast

Genus Lithophyllon

No information available.

Genus Podabacia

Table 64. Sexuality and Reproduction Mode of Podabacia crustacea.
Taxon Sexuality Reproduction
Podabacia crustacea Gonochoric Broadcast

Genus Polyphyllia

Table 65. Sexuality and Reproduction Mode of Polyphyllia talpina.
Taxon Sexuality Reproduction
Polyphyllia talpina Gonochoric Broadcast

Genus Sandalolitha

Table 66. Sexuality and Reproduction Mode of Sandalolitha robusta.
Taxon Sexuality Reproduction
Sandalolitha robusta Gonochoric Broadcast

Genus Verilliofungia

No information available.

Genus Zoopilus

No information available.

Family Meandrinidae

This family has 4 genera found only in the Atlantic. For the purist, the Meandrinidae is sometimes spelled differently from the way it is listed in Veron's Corals of the World (2000).

Genus Ctenella

No information available on the reproduction habits of the one known species (C. chagius) from the western Pacific. This coral has a limited distribution (Veron, 2000).

Genus Dendrogyra (Pillar Coral)

Only one known species (D. cylindrus), an uncommon coral from the Caribbean.

image061.jpg

Figure 32. This Dendrogyra is reproducing asexually via 'budding' but it is also a gonochoric broadcast spawner. Photo courtesy of Julian Sprung.

Table 67. Sexuality and Reproduction Mode of Dendrogyra cylindrus.
Taxon Sexuality Reproduction
Dendrogyra cylindrus Gonochoric Broadcast

Genus Dichocoenia (Caribbean Starlet Coral)

Dichocoenia stokesi has been shown to spawn in September and October of each year (Hoke et al., 2002).

Table 68. Sexuality and Reproduction Mode of Dichocoenia Species.
Taxon Sexuality Reproduction
Dichocoenia stellaris Brooder
Dichocoenia stokesi Gonochoric & Hermaphroditic Broadcast

Genus Eusmilia(Flower Coral)

A beautiful Caribbean coral which makes it a rarity in captivity.

Table 69. Reproduction mode ofEusmilia fastigata.
Taxon Sexuality Reproduction
Eusmilia fastigata Broadcast

Genus Gyrosmilia

Gyrosmilia has a relatively small geographical range and is found only in the West Indian Ocean & Red Sea. No information available on the one species known.

Genus Meandrina (Tan Brain Coral)

Table 70. Reproduction Mode of Two Meandrina Species.
Taxon Sexuality Reproduction
Meandrina areolata ? Brooder
Meandrina meandrites Brooder

Genus Montigyra

No information available on the one species known.

Family Merulinidae

Veron (1986) reports these corals are hermaphroditic broadcast spawners.

Genus Boninastrea

No specific information is available on reproductive habits.

Genus Hydnophora

Table 71. Sexuality and Reproduction Modes of Two Hydnophora Species.
Taxon Sexuality Reproduction
Hydnophora exesa Hermaphroditic Broadcast
Hydnophora rigida Hermaphroditic Broadcast

Genus Merulina

Table 72. Sexuality and Reproduction Modes of Two Merulina Species.
Taxon Sexuality Reproduction
Merulina ampliata Hermaphroditic Broadcast
Merulina scabricula Hermaphroditic Broadcast

Genus Paraclavarina

Paraclavarina specimens are occasionally seen in the trade. Although usually colored tan, their distinct shape makes them a worthwhile addition to a SPS tank. Table 61 uses the outdated name of Clavarina, per Babcock and Heyward's 1986 description.

Table 73. Sexuality and Reproduction Mode of Paraclavarina (Clavarina) triangulis.
Taxon Sexuality Reproduction
Clavarina triangularis Hermaphroditic Broadcast

Genus Scapophyllia

Table 74. Reproduction Mode of a Scapophyllia Species.
Taxon Sexuality Reproduction
Scapophyllia cylindrica Hermaphroditic Broadcast

Family Micrabaciidae

Genus Stephanophyllia

Table 75. Reproduction Mode of a Stephanophyllia Species.
Taxon Sexuality Reproduction
Stephanophyllia formosissima Brooder

Family Mussidae

Genus Acanthastrea

Recent imports from Australia's Great Barrier Reef have included a number of spectacular Acanthastrea specimens. Their fluorescent pigments are showcased in reef aquaria and, not surprisingly, are popular and command high prices - making them a good candidate for captive propagation efforts.

image063.jpg

Figure 33. Imagine a tank full of settled Acanthastrea 'spats'. Photo courtesy of Steve Ruddy.

Table 76. Sexuality and Reproduction Modes of Acanthastrea Species.
Taxon Sexuality Reproduction
Acanthastrea hillae Hermaphroditic Broadcast
Acanthastrea echinata Hermaphroditic Broadcast
Acanthastrea lordhowensis Hermaphroditic Broadcast

Genus Micromussa

Veron (2000) created this genus to distinguish corals containing certain skeletal details from those previously listed as Acanthastrea, and states that these corals are rare. His book lists 3 species. To my knowledge, there is spawning information on only one species.

image065.jpg

Figure 34. The reproductive habits of Micromussa species have only recently been described. Photo courtesy Jake Adams and www.coralidea.com

Table 77. Reproduction Mode of a Micromussa Species.
Taxon Sexuality Reproduction
Micromussa amakuensis Broadcast

Genus Mussismilia

Table 78. Reproduction Mode of Mussismilia Species.
Taxon Sexuality Reproduction
Mussismilia braziliensis Hermaphroditic Broadcast
Mussismilia hartii Hermaphroditic Broadcast
Mussismilia hispida Hermaphroditic Broadcast
Mussismilia sp. Hermaphroditic

Genus Mussa(Flower Coral)

Table 79. Sexuality and Reproduction Mode of Mussa Species.
Taxon Sexuality Reproduction
Mussa sp. Hermaphroditic
Mussa angulosa Brooder

Genus Isophyllia(Stalked Cactus Coral)

Table 80. Sexuality and Reproduction Modes of Isophyllia Species.
Taxon Sexuality Reproduction
Isophyllia dipsacea Gonochoric Brooder (external)
Isophyllia sinuosa Gonochoric Brooder
Isophyllia sp. Gonochoric (?) Broadcast

Isophyllia sinuosa Puberty/Adult Colony Size

Adult I. sinuosa specimens are about 6 inches in diameter.

Genus Isophyllastrea (Rough Star Coral)

Veron (2000) states this coral's proper name as Isophyllia rigida, but I have listed it as the Coral Reef Task Force originally described it.

Table 81. Reproduction Mode of Isophyllastrea/Isophyllia rigida.
Taxon Sexuality Reproduction
Isophyllastrea rigida ? Brooder

Genus Mycetophyllia (Large Cactus Coral)

Table 82. Sexuality and Reproduction Mode of Mycetophyllia Species.
Taxon Sexuality Reproduction
Mycetophyllia ailiciae Hermaphroditic Brooder
Mycetophyllia danaana Brooder
Mycetophyllia ferox Hermaphroditic Brooder
Mycetophyllia lamarckiana Brooder
Mycetophyllia reesi Brooder

Genus Australomussa

No information available.

Genus Blastomussa

No information available.

Genus Cynarina

Table 83. Sexuality and Reproduction Mode of Cynarina lacrymalis.
Taxon Sexuality Reproduction
Cynarina lacrymalis Hermaphroditic Broadcast

Genus Lobophyllia

Table 84. Sexuality and Reproduction Mode of Lobophyllia Species.
Taxon Sexuality Reproduction
Lobophyllia corymbosa Hermaphroditic Broadcast
Lobophyllia hemprichii Hermaphroditic Broadcast
Lobophyllia pachysepta Hermaphroditic Broadcast
Lobophyllia sp. Hermaphroditic

Genus Scolymia (Fungus Coral)

image067.jpg

Figure 35. A strikingly beautiful Scolymia specimen. Photo courtesy Jake Adams and www.coralidea.com

Table 85. Reproduction Mode of Scolymia Species.
Taxon Sexuality Reproduction
Scolymia sp. (Atlantic) ? Brooder
Scolymia cubensis See Scolymia wellsi
Scolymia vitiensis Hermaphroditic Broadcast
Scolymia wellsi Hermaphroditic Brooder
Scolymia wellsi Gonochoric (?) Brooder

Genus Symphyllia

Table 86. Sexuality and Reproduction Mode of Symphyllia Species.
Taxon Sexuality Reproduction
Symphyllia radians Hermaphroditic Broadcast
Symphyllia recta Hermaphroditic Broadcast
Symphyllia sp. Hermaphroditic

Family Oculinidae

Genus Acrhelia

This genus was discontinued by Veron (2000), who considers Acrhelia horrescens to now be Galaxea horrescens. I include this information to avoid confusion.

It does not change the fact that Galaxea/Acrhelia horrescens is a brooder (Kawaguti, 1941).

Table 87. Reproduction Mode of Acrhelia horrescens.
Taxon Sexuality Reproduction
Acrhelia horrescens ? Brooder

Genus Galaxea (Galaxy Corals)

Table 88. Sexuality and Reproduction Mode of Galaxea Species.
Taxon Sexuality Reproduction
Galaxea aspera ? Brooder
Galaxea astreata Hermaphroditic Broadcast
Galaxea fascicularis Hermaphroditic, female sterile Broadcast
Galaxea fascicularis Gonochoric Broadcast
Galaxea horrescens Brooder

Genus Neohelia

Table 89. Sexuality of Neohelia porcellania.
Taxon Sexuality Reproduction
Neohelia porcellana Gonochoric (?)

Genus Oculina (Bush Corals or Ivory Tree Coral)

Table 90. Sexuality and Reproduction Modes of Oculina Species.
Taxon Sexuality Reproduction
Oculina sp. Broadcast
Oculina varicosa Gonochoric Broadcast
Oculina patagonica Gonochoric Broadcast

Oculina varicosa spawns in late summer and early fall. Planula larvae settle at about 21 days of age (Brooke et al., 2002).

Genus Madepora

Table 91. Sexuality of a Madrepora Species.
Taxon Sexuality Reproduction
Madrepora oculata (deep water) Gonochoric ?

Genus Schizoculina

No information available on reproductive habits.

Genus Simplastrea

No information available on reproductive habits.

Family Pectiniidae

All corals in Pectiniidae are hermaphroditic broadcast spawners (Veron, 1986).

Genus Echinomorpha

Contains one species, Echinomorpha (formerly Echinophyllia) nihihirai (Veron, 2000).

Genus Echinophyllia

Some Echinophyllia species are particularly colorful and command extreme prices. Unfortunately, observations of spawning within aquaria have not been made.

image069.jpg

Figure 36. An Echinophyllia species. Photo by the author.

Table 92. Sexuality and Reproduction Modes of Echinophyllia Species.
Taxon Sexuality Reproduction
Echinophyllia aspera Hermaphroditic Broadcast
Echinophyllia orpheensis Hermaphroditic Broadcast

Genus Mycedium

Table 93. Sexuality and Reproduction Mode of a Mycedium Species.
Taxon Sexuality Reproduction
Mycedium elephantotus Hermaphroditic Broadcast

Genus Oxypora

Table 94. Sexuality and Reproduction Modes of Two Oxypora Species.
Taxon Sexuality Reproduction
Oxypora glabra Hermaphroditic Broadcast
Oxypora lacera Hermaphroditic Broadcast

Genus Pectinia

Table 95. Sexuality and Reproduction Modes of Pectinia Species.
Taxon Sexuality Reproduction
Pectinia alcicornis Hermaphroditic Broadcast
Pectinia lactuca Hermaphroditic Broadcast
Pectinia paeonia Hermaphroditic Broadcast

Family Pocilloporidae

Family Pocilloporidae contains 3 genera - Pocillopora, Seriatopora and Stylophora.

Genus Pocillopora (Wart Corals)

image071.jpg

Figure 37. Pocillopora eydouxi is a hermaphroditic broadcast spawner. Kauai, Hawaii. Photo by the author.

Table 96. Sexuality and Reproduction Mode of Pocillopora Species.
Taxon Sexuality Reproduction
Pocillopora bulbosa See P. damicornis Brooder
Pocillopora brevicornis See P. damicornis
Pocillopora caespitosa See P. damicornis Brooder
Pocillopora damicornis Hermaphroditic Brooder
Pocillopora damicornis Parthenogenic?
Pocillopora damicornis Broadcast
Pocillopora damicornis Type Y Brooder
Pocillopora damicornis Type B Brooder
Pocillopora elegans ? Brooder
Pocillopora eydouxi Hermaphroditic Broadcast
Pocillopora meandrina Hermaphroditic Broadcast
Pocillopora verrucosa Hermaphroditic Broadcast
Pocillopora verrucosa ? Broadcast
image073.jpg

Figure 38. In this photo by David Kearnes, the normally clear waters off Kailua-Kona, Hawaii become cloudy with gametes during the seasonal Pocillopora meandrina spawnings.

image075.jpg

Figure 39. Within hours, the water will swarm with billions of planula larvae. Only a few will survive long enough to settle. Photomicrograph by the author.

image078.jpg

Figure 40. Again, this figure demonstrates that a brooder (P. damicornis) is reproductive at a smaller size than the broadcast spawner (P. meandrina).

Puberty Sizes of Two Pocillopora Species

Pocillopora damicornis is probably the most researched species of any stony coral.

Jokiel (1997) states that Pocillopora damicornis at a size of 10 cm (~4") diameter releases 220 planula larvae per month. Harrigan (1972) found P. damicornis as small as 5 cm in diameter can produce planulae (estimated to be 1-2 years old).

Ricmond (in Richmond and Hunter, 1990) states Pocillopora damicornis takes at least two years to mature. P. damicornis colonies in the eastern Pacific can live to at least 70 years of age (Richmond & Hunter, 1990).

During the 2008 Pocillopora meandrina mass spawning in Hawaii, the smallest colony observed spawning was ~6" in diameter (personal observations), which is about 2" less in diameter than the smallest mature colony observed by Stimson (1978).

Genus Seriatopora (Bird's Nest Corals)

image079.jpg

Figure 41. The red morph of Seriatopora hystrix. It is a hermaphroditic brooder, making it a prime candidate for aquarium spawning, yet reports of reproduction in captivity are practically non-existent. Why? (This photo was taken during a recent visit to Salt Lake City. If this is your coral, please contact me and I'll post a tardy photo credit).

Table 97. Sexuality and Reproduction Mode of Seriatopora Species.
Taxon Sexuality Reproduction
Seriatopora caliendrum Hermpahroditic Brooder
Seriatopora hystrix Hermaphroditic Brooder

Genus Stylophora (Club Coral)

Table 98. Sexuality and Reproduction Mode of Stylophora Species.
Taxon Sexuality Reproduction
Stylophora mordax Brooder
Stylophora pistillata Hermaphroditic Brooder
Stylophora pistillata Protandrous Hermaphrodite

Notes on Stylophora pistillata Sexuality

Stylophora pistillata is a male in its first year and then becomes hermaphroditic, capable of self-fertilization and brooding in its second year (Loya, 1976). Later, Rinkevich and Loya stated that Stylophora pistillata reaches puberty at 1.5 to 2 years of age.

image082.jpg

Figure 42. These two brooding corals mature relatively quickly.

Family Poritidae

Genus Alveopora

Table 99. Sexuality and Reproduction Mode of Alveopora Species.
Taxon Sexuality Reproduction
Alveopora daedalea Hermaphroditic Brooder
Alveopora japonica Hermaphroditic Brooder

Genus Goniopora (Flower Pot Corals)

Table 100. Sexuality and Reproduction Mode of Goniopora Species.
Taxon Sexuality Reproduction
Goniopora columna Gonochoric Broadcast
Goniopora dijboutiensis Gonochoric Broadcast
Goniopora fruticosa Possibly sterile
Goniopora gigas ? Broadcast
Goniopora lobata Gonochoric Broadcast
Goniopora minor Gonochoric Broadcast
Goniopora norfolkensis Gonochoric Broadcast
Goniopora palauensis Broadcast
Goniopora queenslandiae Gonochoric Brooder
Goniopora savignyi Gonochoric Broadcast
Goniopora sp. Gonochoric Broadcast
Goniopora sp. 1 Gonochoric Broadcast
Goniopora tenuidens Gonochoric Broadcast

Genus Porites (Lobe and Finger Corals)

image083.jpg

Figure 43. A Porites lutea, from a shallow tide pool in Keauhou, Hawaii. Photo by the author.

Table 101. Sexuality and Reproduction Mode of Porites Species.
Taxon Sexuality Reproduction
Porites andrewsi Gonochoric Broadcast
Porites annae Broadcast
Porites astreoides Hermaphroditic Brooder
Porites astreoides Hermaphroditic & Female Brooder
Porites astreoides Hermaphrodite & Female/ Male Brooder
Porites attenuata Brooder
Porites australensis Gonochoric Broadcast
Porites branneri Brooder
Porites brighami Gonochoric or Hermaphroditic Brooder
Porites clavaria Gonochoric Brooder
Porites colonensis Brooder
Porites compressa Gonochoric Broadcast
Porites cylindrica Gonochoric Broadcast
Porites divaricata Brooder
Porites evermanni Gonochoric Broadcast
Porites furcata Gonochoric Brooder
Porites haddoni see Porites stephensoni Brooder
Porites heronensis Brooder
Porites lichen Hermaphroditic Brooder
Porites lobata Gonochoric Broadcast
Porites lobata Parthenogenic?
Porites lutea Gonochoric Broadcast
Porites lutea Parthenogenic?
Porites murrayensis Gonochoric Brooder
Porites panamensis Gonochoric Brooder
Porites porites Gonochoric Brooder
Porites porites Hermaphroditic
Porites rus Gonochoric ?
Porites solida Gonochoric Broadcast
Porites stephensoni ? Brooder
image086.jpg

Figure 44. Sex ratios of various Porites species.

image088.jpg

Figure 45. Brooders are in green, Broadcasters in orange. Note that adult size of brooders is always smaller that broadcasting species. Also note that 100 cm diameter may not be the largest possible colony size, it is the uppermost category listed by the researcher (Soong, 1993). For true maximum colony sizes of Porites lobata and P. lutea, see Figure 45. Note that P. brighami is endemic to Hawaii.

image090.jpg

Figure 46. True maximum colony size of various Porites species. Information from Doug Fenner, 2005.

image092.jpg

Figure 47. Size of adult colonies of the Atlantic coral Porites astreoides. It is a brooder.

Notes on Porites Puberty Size and Sex Ratios

Porites andrewsi: 2:1 male/female, with a small percentage (~4%) being hermaphroditic.

Porites astreoides: A Caribbean coral, and within specimens containing gonads - some do not!) are found in percentages of 48% hermaphroditic to 52% female, giving a ratio, of course, ~1 to 1 (Chornesky and Peters, 1987). McGuire (1998) states that Porites astreoides capable of reproduction at 70 cm2 (approx. 14" square).

Porites lobata: Spawning ratios were 1:1 in Hawaii (Mate, 1997); ~1:1 at the Great Barrier Reef, Australia (Kojis and Quinn, 1981) and 1:1 in the eastern Pacific (but 14% of Costa Rican P. lobata colonies were hermaphroditic - Glynn et al., 1994).

Porites lutea: Roughly 1:1 at the Great Barrier Reef, Australia (Kojis and Quinn, 1981).

Harriot (1983) estimates Porites lutea reaches puberty at 4 years of age.

Porites murrayensis: Reproductive at 1.3 - 5.5 cm diameter (adult colonies are <20 cm in diameter) at Australia's Great Barrier Reef (Kojis and Quinn, 1981)

Genus Poritipora

This genus, erected by Veron, 2000, has a limited distribution from the east coast of Africa and northeast to the Indian coast and nothing is known of its reproductive habits.

Genus Stylaraea

This genus contains but one species (S. punctata), and it is small (<15mm across). We could expect its reproductive size to be very small. It refers carbonate biofilms for settlement (Golbuu & Richmond, 2007). Information is available for S. punctata, but I lost the reference and am supremely disappointed that I cannot find that information in a stack of over 500 reference papers.

Subgenus Synaraea

Controversy about the validity of subgenus Synaraea exists (Veron 2000 lists Porites (Synaraea) rus as simply P. rus, but acknowledges the previous classification).

Table 102. Sexuality of Porites (SubgenusSynaraea) rus.
Taxon Sexuality Reproduction
Porites rus Gonochoric ?

Family Rhizangiidae

Genus Astrangia

Table 103. Sexuality and Reproduction Modes of Astrangia Species.
Taxon Sexuality Reproduction
Astrangia danae Gonochoric Broadcast
Astrangia lajollaensis Gonochoric Broadcast
Astrangia poculata See A. danae

Genus Culicia

No information available on this corals' reproductive habits.

Family Siderastreidae

Many studies have been conducted on Siderastrea species from the Atlantic, some of which is listed here.

Genus Anomastraea

No information available on this corals' reproductive habits.

Genus Coscinaraea

Table 104. Sexuality and Reproduction Mode of Coscinarea columna.
Taxon Sexuality Reproduction
Coscinarea columna Gonochoric Broadcast

Genus Horastrea

No information available on this coral's reproductive habits.

Genus Psammocora

Table 105. Sexuality and Reproduction Mode of Psammocora species.
Taxon Sexuality Reproduction
Psammocora contigua Gonochoric Broadcast
Psammocora digitata Gonochoric Broadcast
Psammocora stellata ? Brooder

Genus Pseudosiderastrea

No reproductive information available.

Genus Siderastrea (Starlet Corals)

Legally-collected Caribbean Siderastrea colonies (S. siderea, S. stellata or S. radians) may be available in the future (Michael Janes, personal communication). S. radians is relatively small, and generally considered to be a brooder, making it an excellent candidate for captive propagation efforts.

Table 106. Sexuality and Reproduction Mode of Siderastrea species.
Taxon Sexuality Reproduction
Siderastrea radians Gonochoric Brooder
Siderastrea radians Hermaphroditic Brooder
Siderastrea radians Protandrous Hermaphrodite
Siderastrea siderea Brooder
Siderastrea siderea Gonochoric Broadcast
Siderastrea stellata Brooder
image094.jpg

Figure 48. Siderastrea sizes - note that 100 cm diameter may not be the largest colony size, it is the largest category (50-100 cm) listed by the researcher (Soong, 1993).

image096.jpg

Figure 49. Colony size and its relationship to sexual maturity of Siderastrea radians.

Sex Ratios of Siderastrea radians

Siderastrea radians is found in male/female ratios of 1:1. (Lazar, internet resource, date unknown).

image098.jpg

Figure 50. Males and females are found evenly in this case. It could be different in other areas due to any number of factors, but this has yet to be demonstrated for this coral species.

image100.jpg

Figure 51. Siderastrea species have been carefully studied by a number of researchers, resulting in information such as this.

Table 107. Puberty Size and Age Data for Siderastrea Species.
Taxa Puberty Size Age Reference
Siderastrea radians 2- 4 cm² 1.5 - 2 years Soong, 1992
Siderastrea siderea >100 cm² 7-9 years Soong, 1992

Comments on Siderastrea Reproduction and Longevity

Siderastrea radians is a protogynous hermaphrodite (Duerden, 1902, in Fadlallah, 1983).

Elahi and Edmunds (2007) report that Siderastrea siderea colonies can be at least 125 years old (based on a growth rate of 4mm (~1/6") per year). Siderastrea siderea juveniles are less than 40mm in diameter, and mature colonies are 5.9 - 11.4 cm in diameter.

Family Trachyphylliidae

Genus Trachyphyllia

There are limited reports of this coral broadcast spawning in aquaria (Tyree, 1994).

In Closing

As a matter of convenience (mostly mine), I have chosen to publish the reference list at the close of this series. If you like a copy earlier, please email me (RiddleLabs@aol.com) and I'll send this information in a Word document.

Next time, we'll examine the reproduction habits of soft corals.

Acknowledgements

It would have been much easier to simply list the reproductive information of various stony corals. In fact, the initial draft of this article contained few photos. I felt this made the article a little too dry, so, I went, with hat in hand, to many sources asking for help - and thankfully a number of hobbyists and professional aquarists responded. Many thanks go to Dr. Bruce Carlson (once at the Waikiki Aquarium, but now at the Georgia Aquarium), Mitch (the King) Carl of the Henry Doorly Zoo, Omaha, Nebraska, Jake Adams (www.coralidea.com), Steve Ruddy (www.coralreefecosystems.com), Julian Sprung (www.twolittlefishies.com), master aquarist Tony Vargas, Michael P. Janes (www.aquatouch.com), and all the hobbyists across the country who extended a warm welcome to their homes and allowed me to take measurements and photographs.

figure51.jpg

Figure 51. Suggested records for those observing a coral spawning.

Table 108. Quick and Easy Reference Table
Taxon Sexuality Reproduction Vertical (egg) Horizontal water 85% Reference
Acanthastrea hillae Hermaphroditic Broadcast Babcock et al., 1986
Acanthastrea echinata Hermaphroditic Broadcast Bermas et al., 1992
Acanthastrea lordhowensis Hermaphroditic Broadcast Wilson & Harrison, 1997
Acrhelia horrescens ? Brooder Kawaguti, 1941
Acropora (Isopora) brueggemanni Hermaphroditic Brooder Okubo et al., 2007
Acropora (Isopora) cuneata Hermaphroditic Brooder Bothwell, 1981
Acropora (Isopora) palifera Hermaphroditic Brooder Bothwell, 1981
Acropora (Isopora) togianensis Hermaphroditic Brooder Wallace et al., 2007
Acropora aculeus Hermaphroditic Broadcast Babcock et al., 1986
Acropora acuminata Broadcast Rosser & Gilmour, in press
Acropora anthoceris Hermaphroditic Broadcast see van Oppen Babcock et al., 1986
Acropora aspera Hermaphroditic Broadcast Bothwell, 1981
Acropora austera Hermaphroditic Broadcast Babcock et al., 1986
Acropora caroliniana Broadcast Baird et al., 2000
Acropora cerealis Hermaphroditic Broadcast Babcock et al., 1986
Acropora cervicornis Hermaphroditic Broadcast Vargas-Ángel & Thomas, 2002
Acropora clathrata Hermaphroditic Broadcast Babcock et al., 1986
Acropora corymbosa Hermaphroditic Brooder Stimson, 1978
Acropora cytheria Hermaphroditic Broadcast Wallace et al., 2007
Acropora danai Hermaphroditic Broadcast Dai et al., 1992; Hayashibara et al.,1993
Acropora delicatula Possibly sterile Richmond & Hunter, 1990
Acropora delicatula See Acropora selago Broadcast Wallace et al., 2007
Acropora digitifera Hermaphroditic Broadcast Wallace et al., 2007
Acropora dilitata Hermaphroditic Broadcast Heyward, 1985
Acropora divaricata Hermaphroditic Broadcast Babcock et al., 1986
Acropora donei Hermaphroditic Broadcast Guest et al., 2005; Hayashibara et al.,1993
Acropora elseyi Broadcast Marquis et al., 2005
Acropora eurystoma Hermaphroditic Broadcast Shlesinger & Loya, 1985
Acropora exquisita Hermaphroditic Broadcast Hayashibara et al.,1993
Acropora florida Hermaphroditic Broadcast Babcock et al., 1986
Acropora formosa Hermaphroditic Broadcast Babcock et al., 1986
Acropora gemmifera Hermaphroditic Broadcast Morita et al., 2006
Acropora glauca ? Broadcast Babcock et al., 1986
Acropora grandis Hermaphroditic Broadcast Guest et al., 2002; Babcock et al., 1986
Acropora grandulosa Hermaphroditic Broadcast Wallace, 1985
Acropora hemprichii Hermaphroditic Broadcast Rinkevich and Loya, 1979
Acropora horrida Hermaphroditic Broadcast Wallace, 1985
Acropora humilis Hermaphroditic Broadcast Bothwell, 1981
Acropora humilis ? Brooder Stimson, 1978
Acropora hyacinthus Hermaphroditic Broadcast Wallace et al., 2007
Acropora hystrix Hermaphroditic Broadcast Heyward, 1989
Acropora intermeda Broadcast X Guest et al., 2005
Acropora irregularis Hermaphroditic Broadcast Heyward, 1989; Richmond & Hunter, 1990
Acropora jacquelineae Broadcast Baird et al., 2000
Acropora kimbeensis Baird et al., 2000
Acropora "kosurini-like" Broadcast Rosser & Gilmour, in press
Acropora latistella Hermaphroditic Broadcast Babcock et al., 1986
Acropora lianae Broadcast Guest et al., 2005
Acropora listeri Broadcast Harrison et al., 1984
Acropora longicyathus Hermaphroditic Broadcast Babcock et al., 1986
Acropora loripes Hermaphroditic Broadcast Babcock et al., 1986
Acropora lutkeni Hermaphroditic Broadcast Babcock et al., 1986
Acropora microclados Hermaphroditic Broadcast Guest et al., 2005; Hayashibara et al.,1993
Acropora microphthalma Hermaphroditic Broadcast Babcock et al., 1986
Acropora millepora Hermaphroditic Broadcast Babcock et al., 1986
Acropora monticulosa Hermaphroditic Broadcast Babcock et al., 1986
Acropora muricata Hermaphroditic Broadcast Wallace et al., 2007
Acropora nana Hermaphroditic Broadcast Dai et al., 1992
Acropora nastua Hermaphroditic Broadcast Babcock et al., 1986
Acropora nobilis Hermaphroditic Broadcast Babcock et al., 198
Acropora ocellata Hermaphroditic Broadcast Kenyan, 1992
Acropora palawensis ? Brooder Kawaguti, 1940 in Fadlallah, 1983
Acropora palmata Hermaphroditic Broadcast Miller et al., 2000
Acropora palmerae Broadcast Dai et al., 1992
Acropora papillare Broadcast Rosser & Gilmour, in press
Acropora plumosa Broadcast Baird et al., 2000
Acropora polystoma Broadcast Rosser & Gilmour, in press
Acropora pulchra Hermaphroditic Broadcast Babcock et al., 1986
Acropora robusta Hermaphroditic Broadcast Babcock et al., 1986
Acropora samoensis Hermaphroditic Broadcast Guest et al., 2005; Hayashibara et al.,1993
Acropora sarmentosa Hermaphroditic Broadcast Babcock et al., 1986
Acropora scandens Hermaphroditic Broadcast Babcock et al., 1986
Acropora secale Hermaphroditic Broadcast Babcock et al., 1986
Acropora selago Hermaphroditic Broadcast Babcock et al., 1986
Acropora smithi Hermaphroditic Broadcast Richmond & Hunter, 1990
Acropora solitaryensis Hermaphroditic Broadcast Harrison, 2008
Acropora spathulata Broadcast Baird et al., 2000
Acropora spicifera Broadcast Dai et al., 1992
Acropora squarrosa Hermaphroditic Broadcast Kenyon, 1992
Acropora striata ? Brooder Stimson, 1978
Acropora subulata Broadcast Rosser & Gimour, in press
Acropora surculosa Hermaphroditic Broadcast Kenyon, 1992
Acropora tenuis Hermaphroditic Broadcast Babcock et al., 1986
Acropora tortuosa Hermaphroditic Broadcast Harrison, 2008
Acropora valenciennesi Hermaphroditic Broadcast Hayashibara et al.,1993
Acropora valida Hermaphroditic Broadcast Babcock et al., 1986
Acropora variabilis Hermaproditic Broadcast Shlesinger et al., 1998
Acropora vaughani Broadcast Rosser & Gimour, in press
Acropora verweyi Hermaphroditic Broadcast Guest et al., 2005; Hayashibara et al.,1993
Acropora willisae Broadcast Negri et al., 2001
Acropora yongei Hermaphroditic Broadcast Babcock et al., 1986
Agaricia agaricites Hermaphroditic Brooder Gleason et al., 2001
Agaricia crassa ? Brooder Vaughan, 1908; in Fadlallah, 1983
Agaricia fragilis ? Brooder Mayor, 1915; in Fadlallah, 1983
Agaricia grahame ? Brooder NOAA Data base
Agaricia humilis Hermaphroditic Brooder van Moorsel, 1980
Agaricia lamarcki ? Brooder NOAA Data base
Agaricia purpurea Hermaphroditic Brooder van Moorsel, 1980
Agaricia tenufolia ? Brooder NOAA Data base
Agaricia undata ? Brooder NOAA Data base
Alveopora daedalea Hermaphroditic Brooder Shlesinger et al., 1998
Alveopora japonica Hermaphroditic Brooder Igarashi et al., 1992
Anacropora matthai Broadcast Wallace et al., 2007
Astrangia danae Gonochoric Broadcast Szmant-Froelich et al., 1980
Astrangia lajollaensis Gonochoric Broadcast Fadlallah, 1982
Astrangia poculata See A. danae Fitzgerlad & Szmant, 1997
Astreopora gracilis Broadcast Dai et al., 1992
Astreopora listeri Broadcast Dai et al., 1992
Astreopora myriophthalma Hermaproditic Broadcast Shlesinger et al., 1998
Astreopora randalli Hermaproditic Broadcast Richmond & Hunter, 1990
Australogyra zelli Hermaphroditic Broadcast Babcock et al., 1986
Balanophyllia elegans Gonochoric Brooder Fadlallah & Pearse, 1982
Balanophyllia europaea Hermaphroditic Brooder Goffredo et al., 2005
Balanophyllia pruvoti Gonochoric Radetic et al., 2002
Balanophyllia regia Gonochoric (?) Brooder Goffredo et al., 2005
Balanophyllia sp. Brooder Abe, 1937, in Fadlallah, 1983
Barabattoia amicorum Hermaphroditic Broadcast Babcock et al., 1986
Catalaphyllia jarderni Gonochoric Broadcast Willis et al., 1995
Caulastrea furcata Hermaphroditic Broadcast Babcock et al., 1986
Cladocora caespitosa Hermaphroditic Broadcast Kružić et al., 2008
Cladopsammia rolandi Hermaphroditic Brooder de Lacaze-Duthiers, 1897, in Fadlallah, 1983
Clavarina triangularis Hermaphroditic Broadcast Babcock & Heyward, 1986; Babcock et al., 1986
Colpophyllia amaranthus Broadcast NOAA Database
Colpophyllia breviseralis Broadcast NOAA Database
Colpophyllia natans Hermaphroditic Broadcast NOAA Database
Coscinarea columna Gonochoric Broadcast Babcock et al., 1986
Cynarina lacrymalis Hermaphroditic Broadcast Shlesinger et al., 1998
Cyphastrea chalcidicum Hermaphroditic Broadcast Dai et al., 1992; Babcock et al., 1986
Cyphastrea microphthalma Hermaphroditic Broadcast Shlesinger et al., 1998
Cyphastrea ocellina Hermaphroditic Brooder Stimson, 1978; Kolinski & Cox, 2003
Cyphastrea seraila Hermaphroditic Broadcast Wilson, 1997; Hayashibara et al.,1993
Dendrogyra cylindrus Gonochoric Broadcast Szmant, 1986
Dendrophyllia nigrescens See Tubastraea spp.
Dendrophyllia sp. Gonochoric Brooder Babcock et al., 1986
Diaseris distorta Gonochoric Broadcast Colley et al., 2000
Dichocoenia stellaris Brooder NOAA Data base
Dichocoenia stokesi Gonochoric & Hermaphroditic Broadcast Hoke et al., 2002
Diploastrea heliopora Gonochoric Broadcast Hayashibara et al.,1993
Diploria clivosa Hermaphroditic Broadcast Alvarado et al., 2003; NOAA Data base
Diploria labyrinthiformis Hermaphroditic Broadcast Alvarado et al., 2003
Diploria labyrinthiformis Hermaphroditic Brooder Duerden, 1902 in Fadlallah, 1983
Diploria strigosa Hermaphroditic Broadcast Alvarado et al., 2003
Diploria strigosa Hermaphroditic Brooder Hagman et al., 1998
Echinophyllia aspera Hermaphroditic Broadcast Dai et al., 1992; Fan & Dai, 1992
Echinophyllia orpheensis Hermaphroditic Broadcast Babcock et al., 1986
Echinopora gemmacea Hermaphroditic Broadcast Babcock & Heyward, 1986
Echinopora horrida Hermaphroditic Broadcast Babcock et al., 1986
Echinopora lamellosa Hermaphroditic Broadcast Babcock et al., 1986
Echinopora pacificus Hermaphroditic Broadcast Hayashibara et al.,1993
Enallopsammia rostrata( deep water) Gonochoric Waller, 2005
Euphyllia ancora Gonochoric Broadcast Willis et al., 1985
Euphyllia divisa Gonochoric Broadcast Babcock et al., 1986
Euphyllia glabrescens Hermaphroditic Brooder Huang et al., 2008
Euphyllia parancora Gonochoric? Broadcast ASIRA Data base
Euphyllia rugosa Brooder ASIRA Data base
Eusmilia fastigata Broadcast Coral Reef Task Force Data base
Favia bennettae Hermaphroditic Broadcast Babcock et al., 1986
Favia doreyensis Hermaphroditic Broadcast Marshall & Stephenson, 1933, in Fadlallah, 1983
Favia favus Hermaphroditic Broadcast Shlesinger et al., 1998
Favia favus Gonochoric Broadcast Alvarado et al., 2003
Favia favus Hermaphroditic Broadcast Hayashibara et al.,1993
Favia fragum Hermaphroditic Brooder Alvarado et al., 2003
Favia helianthoides Hermaphroditic Broadcast Bermas et al., 1992
Favia laxa Broadcast Dai et al., 1992
Favia lizardensis Hermaphroditic Broadcast Babcock et al., 1986
Favia matthai Hermaphroditic Broadcast Babcock et al., 1986
Favia pallida Hermaphroditic Broadcast Penland et al., 2004; Babcock & Hey ward,1986
Favia rotumana Broadcast Harrison et al., 1984
Favia speciosa Hermaphroditic Broadcast Penland et al., 2004; Dai et al, 1992
Favia stelligera Hermaphroditic Broadcast Babcock et al., 1986
Favia veroni Hermaphroditic Broadcast Babcock et al., 1986
Favites abdita Hermaphroditic Broadcast Penland et al., 2004; Wallace et al., 2007
Favites bennettae Hermaphroditic Broadcast Babcock et al., 1986
Favites chinensis Hermaphroditic Broadcast Nowzawa & Harrison, 2005
Favites complanata Hermaphroditic Broadcast Babcock & Heyward, 1986
Favites flexuosa Hermaphroditic Broadcast Babcock et al., 1986
Favites halicora Hermaphroditic Broadcast Penland et al., 2004; Babcock et al., 1986
Favites pentagona Hermaphroditic Broadcast Oliver and Babcock, 1992
Favites russelli Hermaphroditic Broadcast Dai et al., 1992; Babcock et al., 1986
Flabellum alabastrum (deep water) Gonochoric Fadlallah, 1983
Flabellum angulare (deep water) Gonochoric Waller, 2005
Flabellum curvatum (deep water) Gonochoric Waller, 2005
Flabellum impensum (deep water) Gonochoric Waller, 2005
Flabellum japonicum Gonochoric Fadlallah, 1983
Flabellum rubrum Protandrous Hermaphrodite Moseley, 1881, in Fadlallah, 1983
Flabellum rubrum Hermaphroditic Fadlallah, 1983
Flabellum thouarsii (deep water) Gonochoric Waller, 2005
Fungia actiniformis See Heliofungia actiniformis
Fungia concinna Gonochoric Broadcast X Oliver, 1985
Fungia fungites Gonochoric Broadcast Penland et al., 2004; Babcock et al., 1986
Fungia granulosa Gonochoric Broadcast Kramarsky-Winter & Loya, 1998
Fungia horrida Broadcast Babcock et al., 2003
Fungia paumotensis Gonochoric Broadcast Willis et al., 1985
Fungia repanda Gonochoric Broadcast Hayashibara et al.,1993
Fungia scutaria Gonochoric Broadcast Kolinski & Cox, 2003
Fungia scutaria Parthenogenic? Krupp, 1983
Fungia sp. (Okinawa) Gonochoric Broadcast Heyward et al., 1987
Fungicyathus crispa Gonochoric Waller, 2005
Fungicyathus fragilis Gonochoric Waller, 2005
Fungiacyathus marenzelleri Gonochoric Waller, 2005
Galaxea aspera ? Brooder Fadlallah, 1983
Galaxea astreata Hermaphroditic Broadcast Dai et al., 1992; Babcock et al., 1986
Galaxea fascicularis Hermaphroditic, female sterile Broadcast Shlesinger et al., 1998; Hayakawa et al., 2005
Galaxea fascicularis Gonochoric Broadcast Harrison & Wallace, 1990; Hayakawa et al., 2005
Galaxea horrescens Brooder ASIRA Data base
Gardineroseris planulata Hermaphroditic Glynn et al., 1996
Goniastrea aspera Hermaphroditic Broadcast Penland et al., 2004
Goniastrea aspera Hermaphroditic Brooder Nishikawa & Sakai, 2003
Goniastrea australensis Hermaphroditic Broadcast Penland et al., 2004; Wilson & Harrison, 2005
Goniastrea edwardsi Hermaphroditic Broadcast Penland et al., 2004
Goniastrea favulus Hermaphroditic Broadcast X Alvarado et al., 2003
Goniastrea favulus Protandrous Hermaphrodite Kojis & Quinn, 1981
Goniastrea palauensis Hermaphroditic Broadcast Penland et al., 2004
Goniastrea pectinata Hermaphroditic Broadcast Penland et al., 2004
Goniastrea retiformis Hermaphroditic Broadcast Shlesinger et al., 1998
Goniocorella dumosa (deep water) Gonochoric Waller, 2005
Goniopora columna Gonochoric Broadcast Willis et al., 1985
Goniopora dijboutiensis Gonochoric Broadcast Willis et al., 1985
Goniopora fruticosa Possibly sterile Richmond & Hunter, 1990
Goniopora gigas ? Broadcast Petersen et al., 2007
Goniopora lobata Gonochoric Broadcast Babcock & Heyward, 1986; Babcock et al., 1986
Goniopora minor Gonochoric Broadcast Babcock et al., 1986
Goniopora norfolkensis Gonochoric Broadcast Babcock et al., 1986
Goniopora palauensis Broadcast Babcock et al., 1986
Goniopora queenslandiae Gonochoric Brooder Yamazato et al., 1975, in Fadlallah, 1983
Goniopora savignyi Gonochoric Broadcast Shlesinger et al., 1998
Goniopora sp.  Gonochoric Broadcast Babcock et al., 1986
Goniopora sp. 1 Gonochoric Broadcast Babcock et al., 1986
Goniopora tenuidens Gonochoric Broadcast Willis et al., 1985
Heliofungia actiniformis Hermaphroditic (?) Brooder Abe, 1937 in Krupp, 1983; Fadlallah, 1983
Heliofungia actiniformis Gonochoric Broadcast Willis et al., 1985;  Babcock et al., 1986
Helioseris cucullata Brooder NOAA Data base
Herpetoglossa simplex Gonochoric Broadcast Babcock et al., 1986
Herpolitha limax Gonochoric Broadcast Babcock et al., 1986
Heteropsammia aequicostatus Gonochoric Broadcast Fisk, 1981
Heteropsammia cochlea Gonochoric Broadcast Fisk, 1981
Hydnophora exesa Hermaphroditic Broadcast Babcock et al., 1986
Hydnophora rigida Hermaphroditic Broadcast Babcock et al., 1986
Isophyllastrea rigida ? Brooder Coral Reef Task Force data
Isophyllia dipsacea Gonochoric Brooder (external) Fadlallah, 1985
Isophyllia sinuosa Gonochoric Brooder Fadlallah, 1983
Isophyllia sp. Gonochoric (?) Broadcast Babcock et al., 1986
Isopora brueggemanni Hermaphroditic Brooder  Okubo et al., 2007
Isopora togianensis Hermaphroditic Brooder Wallace et al., 2007
Leptastrea bottae Gonochoric Broadcast Kolinski & Cox, 2003
Leptastrea purpurea Gonochoric Broadcast Kolinski & Cox, 2003
Leptopsammia pruvoti Gonochoric Brooder X Kružić et al., 2008
Leptoria phryia Hermaphroditic Broadcast Dai et al., 1992; Babcock et al., 1986
Lobophyllia corymbosa Hermaphroditic Broadcast Harriot, 1983
Lobophyllia hemprichii Hermaphroditic Broadcast Willis et al., 1985
Lobophyllia pachysepta Hermaphroditic Broadcast Marquis et al., 2005; Babcock et al., 1986
Lobophyllia sp. Hermaphroditic Marshall & Stephenson, 1933; in Fadlallah, 1983
Lophelia pertusa (deep water) Gonochoric ? Waller, 2005; Waller & Tyler, 2005
Madracis carmabi Hermaphroditc Brooder Vermeij et al., 2004
Madracis decactis Hermaphroditc Brooder Vermeij et al., 2004
Madracis formosa Hermaphroditc Brooder Vermeij et al., 2004
Madracis mirabilis Hermaphroditc Brooder Vermeij et al., 2004
Madracis pharensis Hermaphroditc Brooder X Vermeij et al., 2004
Madracis senaria Hermaphroditc Brooder X Vermeij et al., 2004
Madrepora oculata (deep water) Gonochoric ? Waller & Tyler, 2005
Manicina areolata Hermaphroditic Brooder Fadlallah, 1983
Meandrina areolata ? Brooder Yonge, 1935, in Fadlallah, 1983
Meandrina meandrites Brooder NOAA Data base
Merulina ampliata Hermaphroditic Broadcast Penland et al., 2004; Babcock et al., 1986
Merulina scabricula Hermaphroditic Broadcast Hayashibara et al.,1993
Micromussa amakuensis Broadcast Mezaki et al., 2007
Montastrea  magnistellata Hermaphroditic Broadcast Babcock et al., 1986
Montastrea annularis Hermaphroditic Broadcast Alvarado et al., 2003
Montastrea annuligera Hermaphroditic Broadcast Babcock et al., 1986
Montastrea cavernosa Gonochoric Broadcast Alvarado et al., 2003
Montastrea curta Hermaphroditic Broadcast Wilson, 1997; Babcock et al., 1986
Montastrea faveolata Hermaphroditic Broadcast X Levitan et al., 2004; Steiner, 1995
Montastrea franksi Hermaphroditic Broadcast Levitan et al., 2004; Szmant et al., 1997
Montastrea valenciennesi Hermaphroditic Broadcast Dai et al., 1992; Babcock et al., 1986
Montipora aequituberculata Hermaphroditic Broadcast Dai et al., 1992
Montipora altasepta Hermaphroditic Broadcast X Penland et al., 2004
Montipora cactus Hermaproditic Broadcast Penland et al., 2004; Wallace et al., 2007
Montipora capitata Hermaproditic Broadcast Heyward, 1986
Montipora crassituberculata Hermaproditic Penland et al., 2004
Montipora digitata Hermaproditic Broadcast Penland et al., 2004; Morita et al., 2006
Montipora dilatata Hermaproditic Broadcast Heyward, 1986
Montipora efflorescens Hermaphroditic Broadcast Wallace et al., 2007
Montipora effusa Broadcast Yeemin, 1988
Montipora erythraea Hermaproditic Broadcast Shlesinger et al., 1998
Montipora eydouxi Hermaphroditic Broadcast Hirose et al., 2000
Montipora faveolata Hermaphroditic Broadcast Richmond & Hunter, 1990
Montipora flabellata Hermaphroditic Broadcast Heyward, 1986
Montipora floweri Hermaphroditic Broadcast Hayashibara et al.,1993
Montipora foliosa Hermaphroditic Broadcast n/a n/a Dai et al., 1992
Montipora hispida Hermaproditic Broadcast Penland et al., 2004; Babcock et al., 1986
Montipora informis Hermaphroditic Broadcast Dai et al., 1992
Montipora monasteriata Hermaphroditic Broadcast Babcock et al., 1986
Montipora patula Hermaphroditic Broadcast Heyward, 1986
Montipora peltiformis Broadcast Mundy & Babcock, 1998
Montipora ramosa Broadcast Harrison et al., 1984
Montipora samarensis Hermaphroditic Penland et al., 2004
Montipora spumosa Hermaphroditic Broadcast Babcock et al., 1986
Montipora studeri Hermaphroditic Broadcast Heyward, 1986
Montipora tuberculosa Broadcast Babcock & Heyward, 1986
Montipora turgescens Hermaphroditic Broadcast Babcock et al., 1986
Montipora turtlenesis Hermaphroditic Broadcast Babcock et al., 1987
Montipora venosa Hermaphroditic Broadcast Dai et al., 1992; Hayashibara et al.,1993
Montipora verrilli Hermaphroditic Broadcast Heyward, 1986
Montipora verrucosa Hermaphroditic Broadcast Maté et al., 1998
Moseleya latistellata Hermaphroditic Brooder Wallace et al., 2007
Mussa sp. Hermaphroditic Steiner, 1993
Mussa angulosa Brooder Coral Reef Task Force data
Mussismilia braziliensis Hermaphroditic Broadcast Pires et al., 1999
Mussismilia hartii Hermaphroditic Broadcast Pires et al., 2002
Mussismilia hispida Hermaphroditic Broadcast Pires et al., 2002
Mussismilia sp. Hermaphroditic Pires et al., 1999
Mycedium elephantotus Hermaphroditic Broadcast Babcock & Heyward, 1986
Mycetophyllia ailiciae Hermaphroditic Brooder Szmant, 1986
Mycetophyllia danaana Brooder Coral Reef Task Force data
Mycetophyllia ferox Hermaphroditic Brooder Szmant, 1986
Mycetophyllia lamarckiana Brooder Coral Reef Task Force data
Mycetophyllia reesi Brooder X Coral Reef Task Force data
Oulastrea crispa Hermaphroditic Broadcast Lam, 2000
Oulastrea crispa Hermaphroditic Brooder Lam, 2000
Neohelia porcellana Gonochoric (?) Pratt, 1900, in Fadlallah, 1983
Oculina sp. Broadcast Brooke & Young, 2003
Oculina varicosa Gonochoric Broadcast Brooke & Young, 2003
Oculina patagonica Gonochoric Broadcast Fine et al., 2001
Oulophyllia bennettae Hermaphroditic Broadcast Bermas et al., 1992
Oulophyllia crispa Hermaphroditic Broadcast Babcock et al., 1986
Oxypora glabra Hermaphroditic Broadcast Babcock et al., 1986
Oxypora lacera Hermaphroditic Broadcast Mundy & Babcock, 1998; Babcock et al., 1986
Pachyseris rugosa Gonochoric Broadcast Dai et al., 1992; Babcock et al., 1986
Pachyseris speciosa Gonochoric Broadcast Penland et al., 2004; Baird et al., 2001
Pavona cactus Gonochoric ? Marshall & Stephenson, 1933; in Fadlallah, 1983
Pavona duerdeni Gonochoric Broadcast Kolinski & Cox, 2003
Pavona explanata Gonochoric (?) Stimson, 1978
Pavona gigantea Gonochoric Broadcast Glynn et al., 1996
Pavona varians Gonochoric Broadcast Maté, 1998; Shlesinger et al., 1998
Pectinia alcicornis Hermaphroditic Broadcast Penland et al., 2004; Babcock & Hey ward,1986
Pectinia lactuca Hermaphroditic Broadcast Bermas et al., 1992
Pectinia paeonia Hermaphroditic Broadcast Babcock & Heyward, 1986
Physogyra lichtensteini Gonochoric Broadcast Babcock et al., 1986 & 2003
Platygyra contorta Hermaphroditic Broadcast Hayashibara et al.,1993
Platygyra daedalea Hermaphroditic Broadcast Mangubhai & Harrison, 2008
Platygyra lamellina Hermaphroditic Broadcast Shlesinger et al., 1998; Dai et al., 1992
Platygyra pini Hermaphroditic Broadcast Mangubhai & Harrison, 2008
Platygyra ryukuensis Hermaphroditic Broadcast Heyward, 1988
Platygyra sinensis Hermaphroditic Broadcast Wallace et al., 2007; Oliver and Babcock, 1992
Platygyra verweyi Hermaphroditic Broadcast Mangubhai & Harrison, 2008
Plerogyra sinuosa Broadcast ASIRA Data base
Plesiastrea verispora Broadcast Dai et al., 1992
Pocillopora bulbosa See P. damicornis Brooder Goffredo et al., 2006
Pocillpora brevicornis See P. damicornis
Pocillopora caespitosa See P. damicornis Brooder X Benayahu et al., 1992
Pocillopora damicornis Hermaphroditic Brooder Ward, 1992
Pocillopora damicornis Parthenogenic? Permata et al., 2000
Pocillopora damicornis Broadcast Ward, 1992
Pocillopora damicornis Type Y Brooder Stimson, 1987
Pocillopora damicornis Type B Brooder Stimson, 1987
Pocillopora elegans ? Brooder Stimson, 1978
Pocillopora eydouxi Hermaphroditic Broadcast Hirose et al., 2000
Pocillopora meandrina Hermaphroditic Broadcast Stimson, 1978
Pocillopora verrucosa Hermaphroditic Broadcast Shlesinger et al., 1998
Pocillopora verrucosa ? Broadcast Stimson, 1978
Podabacia crustacea Gonochoric Broadcast Babcock et al., 1986
Polyphyllia talpina Gonochoric Broadcast Willis et al., 1985
Porites andrewsi Gonochoric Broadcast Kojis & Quinn, 1981
Porites annae Broadcast Dai et al., 1992
Porites astreoides Hermaphroditic Brooder Brazeau et al., 1998
Porites astreoides Hermaphroditic & Female Brooder Chornesky & Peters, 1987
Porites astreoides Hermaphrodite & Female/ Male Brooder Soong, 1991
Porites attenuata Brooder ASIRA Data base
Porites australensis Gonochoric Broadcast Neves, 1998
Porites branneri Brooder X Coral Reef Task Force data
Porites brighami Gonochoric or Hermaphroditic Brooder X Hunter & Hodgson in Richmond & Hunter, 1990
Porites clavaria Gonochoric Brooder Duerden, 1902, in Fadlallah, 1983
Porites colonensis Brooder Coral Reef Task Force data
Porites compressa Gonochoric Broadcast Kolinski & Cox, 2003
Porites cylindrica Gonochoric Broadcast Babcock et al., 1986
Porites divaricata Brooder Coral Reef Task Force Data
Porites evermanni Gonochoric Broadcast Neves, 1998
Porites furcata Gonochoric Brooder Soong, 1991
Porites haddoni see Porites stephensoni Brooder Soong, 1991
Porites heronensis Brooder (?) Harriot & Banks, 1995
Porites lichen Hermaphroditic Brooder Kolinski & Cox, 2003
Porites lobata Gonochoric Broadcast Penland et al., 2004
Porites lobata Parthenogenic? Fadlallah, 1983
Porites lutea Gonochoric Broadcast Richmond & Hunter, 1990
Porites lutea Parthenogenic? Fadlallah, 1983
Porites murrayensis Gonochoric Brooder Neves, 1998
Porites panamensis Gonochoric Brooder Glynn et al., 2008
Porites porites Gonochoric Brooder Tomascik & Sander, 1987
Porites porites Hermaphroditic Tomascik & Sander, 1987
Porites rus Gonochoric ? Richmond & Hunter, 1990
Porites solida Gonochoric Broadcast Babcock et al., 1986
Porites stephensoni ? Brooder Soong, 1993
Psammocora contigua Gonochoric Broadcast Babcock et al., 1986
Psammocora digitata Gonochoric Broadcast Babcock et al., 1986
Psammocora stellata ? Brooder Kolinski & Cox, 2003
Rhizopsammia minuta Brooder Abe, 1939, in Fadlallah, 1983
Sandalolitha robusta Gonochoric Broadcast Babcock et al., 1986
Scapophyllia cylindrica Hemaphroditic Broadcast X Willis et al., 1985
Scolymia sp. (Atlantic) ? Brooder Coral Reef Task Force Data base
Scolymia cubensis See Scolymia wellsi
Scolymia vitiensis Hemaphroditic Broadcast Willis et al., 1985
Scolymia wellsi Hemaphroditic Brooder X Pires et al., 2000
Scolymia wellsi Gonochoric (?) Brooder Pitombo, 1992
Seriatopora caliendrum Hermpahroditic Brooder Shlesinger et al., 1998
Seriatopora hystrix Hermpahroditic Brooder Fadlallah, 1983
Siderastrea radians Gonochoric Brooder Soong, 1991
Siderastrea radians Hermaphroditic Brooder X Duerden, 1902, in Coral Reef Task Force Data base
Siderastrea radians Protandrous Hermaphrodite Duerden, 1902, in Fadlallah, 1983
Siderastrea siderea Brooder Neves et al., 2008
Siderastrea siderea Gonochoric Broadcast Szmant, 1986
Siderastrea stellata Brooder Neves et al., 2008
Solenastrea boumoni Broadcast Coral Reef Task Force Data base
Solenastrea hyades Broadcast Coral Reef Task Force Data base
Solensomilia variabilis Gonochoric Broadcast (?) NOAA Technical Memo NMFS-OPR-28
Stephanocoenia intercepta Gonochoric Broadcast Coral Reef Task Force data
Stephanocoenia michelini Gonochoric Broadcast Smith, 1997
Stephanophyllia formosissima Brooder Moseley, 1881, in Fadlallah, 1983
Stylocoeniella sp. Gonochoric Harrison, 1985
Stylophora mordax Brooder Brown et al., 1998
Stylophora pistillata Hermaphroditic Brooder Shlesinger & Loya, 1985
Stylophora pistillata Protandrous Hermaphrodite Loya, 1976
Symphyllia radians Hermaphroditic Broadcast Babcock et al., 1986
Symphyllia recta Hermaphroditic Broadcast Babcock et al., 1986
Symphyllia sp. Hermaphroditic Willis et al., 1985
Tubastraea aurea See Tubastraea coccinea
Tubastraea coccinea Hermaphroditic Brooder Paz-Garcia et al., 2007
Tubastraea faulkneri Gonochoric Brooder Babcock et al., 1986
Turbinaria frondens Gonochoric Willis et al., 1985
Turbinaria mesenterina Broadcast Willis, 1987
Turbinaria reniformis Gonochoric Broadcast X Willis et al., 1985
Turbinaria sp. Gonochoric Broadcast Harrison et al., 1984


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