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.

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.

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)

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).

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.

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)

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)

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).

Genus Turbinaria (Pagoda Corals)

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

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)

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

Maximum Colony Sizes of Brooding and Broadcasting Euphyllia Species

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.

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.

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).

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.

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.

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

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.

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).

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

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.

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

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

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

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

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

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.

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.

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.

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)

Vero