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Blastomussa wellsi, (Wijsman-Best 1973)


Caryl-Ann Jun Ng 2018

Summary

Blastomussa wellsi is a stony coral that is commonly known as Blastomussa Pineapple Coral or the Big Polyp Blastomussa Coral. It is a popular marine aquarium animal and is often mistaken for mushroom or brain corals. Aside from the morphological distinction between the members of Blastomussa, there is a scarce amount of existing research of this genus, which might owe to their rare occurrences in their geographical ranges. Apart from their morphology, aspects such as their life history and behaviour are unclear and therefore will be based on published literature of similar corals of the same order (scleractinia). The classification of the Blastomussa genus into a family is currently under debate, with their current placement in Plesiastreidae or incertae sedis. The current status of B. wellsi on the IUCN Red List is Near Threatened, and the biggest threats they face are commercial harvesting for the aquarium trade and climate change. However, there are no known conservation efforts for this species.

In this study, for the acquisition of a more comprehensive understanding of the mysterious species, specimens of B. wellsi were examined to determine if they fluoresced naturally. The results are outlined in Fluorescence of the Anatomy & Physiology section.


 

Figure 1. B. wellsi polyps of different colour morphs. 



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Figure 1

Physical Description

The Blastomussa genus is distinguished from other colonial members of the Mussidae family that they were initially classified in, by the presence of extratentacular budding from the edge-zone, smaller corallites, and thus proportionally fewer septa (Wells 1968).

Blastomussa wellsi are often fluorescent (see ‘Fluorescence’ in the ‘Anatomy & Physiology’ section), with brightly coloured fleshy polyps ranging from orange to bright red or light brown with rugged mantle vesicles that are occasionally of a lighter colour (Figure 2). The oral disc is sometimes a different colour to the rest of the animal (Benzoni et al 2014), including shades of green, red, or dark grey (Veron & Stafford-Smith 2000). During the day time, the underlying growth form of the species is often concealed by the extended mantles which form a continuous obscuring surface (Veron & Stafford-Smith 2000). In the day, polyp tentacles are also often retracted towards the oral disc (Figures 3 & 4) (Benzoni et al 2014). 


  

Figure 2. (left) Mantle vesicles of a B. wellsi polyp viewed under a light microscope, is of a different colour to the rest of the animal.
Figure 3. (right) Tentacles and mantle vesicles of a B. wellsi polyp viewed under a light microscope. 


The size of each corallite ranges from 8 to 13 millimetres in diameter and are round or oval shaped (Benzoni et al 2014). B. wellsi colonies are usually phaceloid with regularly spaced corallites that rarely show any fusion of corallite walls (Figures 4 & 5), but occasionally colonies are partially-cerioid due to individuals that were budded recently that still maintain connections (Benzoni et al 2014).


  

Figure 4. (top) A phaceloid colony of B. wellsi from the Red Sea. Image source: Australian Institute of Marine Science (2013).
Figure 5. (bottom) B. wellsi corallite arrangement. Image source: Australian Institute of Marine Science (2013). 




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Figure 2
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Figure 3
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Figure 4
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Figure 5

Ecology

Habitat

B. wellsi are found on the outer reef slops of up to 40 meters in depth, living in well-lit, waved-exposed environments (Benzoni et al 2014). According to Benzoni et al (2014), the species recorded at New Caledonia were only observed below a depth of 15 meters.

Predators

Predators and parasites of scleractinian corals are diverse, including but not restricted to gastropods, polychaetes, crustaceans, pycnogonids, asteroids, turtles, and both bony and cartilaginous fishes (Robertson 1970). While the majority of these predators are facultative or obligate parasites, some gastropods and crustaceans live in symbiosis with shallow water, tropical scleractinians (Robertson 1970). As coral larvae are a favoured food resource of many larger filter feeders and fish, their reproduction method must ensure some successful fertilization and the continuance of the population (see ‘Reproduction’ in the ‘Life History & Behaviour’ section). Though apparent damage to the corals by corallivores – consumers of live coral tissue, may seem insignificant, coral fitness and rate of population decline is affected by corallivory (Rotjan et al 2006). Some predator species such as the Crown-of-thorns starfish (Acanthaster planci) are capable of causing extreme damage (Lourey et al 2000; Rotjan & Lewis 2008). No studies have been conducted specifically reviewing predators of Blastomussa spp.

Symbiosis

The brilliant colours of corals are caused by the many unicellular dinoflagellate algae that they host in the gastrodermal or epidermal tissue of their polyps (Schlichter et al 1994) (Figure 6). The mutualistic symbiosis between corals and zooxanthellae allows corals to thrive in nutrient poor waters, with the zooxanthellae providing their host with organic compounds from photosynthesis (glycerol, glucose, and alanine) in exchange for the metabolic wastes of the corals (nitrogen and phosphorus). This relationship and the maintenance of a viable zooxanthellae population dictates if a coral may contribute to the formation of reefs (Brusca et al 2016).


 

Figure 6. The general anatomy of a coral polyp, depicting their symbiotic zooxanthellae in the gastroderm. Adapted from Sabouralt et al (2009). 



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Figure 6

Ecological Importance

Scleractinian corals are ecosystem engineers of tropical reefs, providing the structural framework that supports a plethora of marine species (Jones et al 1994) and protects coastlines (Stat & Gates 2011). As a dominant reef-builder, these corals are capable of creating highly productive habitats of otherwise oligotrophic and barren low-latitudinal, shallow waters (Hoegh-Guldberg 1999). Severe knock-off effects can be anticipated with the decline in populations of reef-building corals such as B. wellsi, such as the decline of populations of other marine species within the reef, and algal phase shifts (Mumby 2009).

Life History and Behaviour

Life Cycle

Due to their rare occurrence in their range (Wilson 2013) and the scarce amount of research conducted on Blastomussa, this section will mostly focus on the life history and behaviour of similar corals of the same order (scleractinia).


Anthozoa is the one class within cnidaria whose members life cycles do not have a medusa stage. Their life cycle is biphasic, with the direct development of a juvenile pelagic planktonic larval phase and a benthic adult phase that follows metamorphosis and settlement (Jones et al 2015).

Figure 7 depicts the typical life cycle of a scleractinian coral, and is the most probable life cycle of B. wellsi. Fertilisation occurs externally or internally, forming a zygote, which then starts to cleavage to form a coeloblastula. Following blastulation, gastrulation occurs to form the endoderm and ectoderm and eventually, the ciliated planula larva (Brusca et al 2016). The planula larva then elongates over several days and swims in the pelagic using its flagella and the ocean currents for a period of days to months (Hayward et al 2011). The planktonic planula larvae then begin metamorphosis with the early morphogenesis of a pharynx, tentacles and septa before larval settlement occurs (Ruppert et al 2009). When the larvae find a suitable substrate and are competent, they settle on their aboral end and metamorphosis continues, where tentacle growth begins on the oral end (Brusca et al 2016). The juvenile polyp begins calcification and symbiotic zooxanthellae is obtained (Hayward et al 2011) before a colony is eventually propagated asexually, thus restarting the cycle.


 

Figure 7. The typical life cycle of a scleractinian coral. Adapted from Brusca et al (2016) and Jones et al (2015). 


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Figure 7

Reproduction

Blastomussa are simultaneous hermaphrodites that utilise both forms of reproduction – sexual and asexual.

One distinguishing character of members of the Blastomussa genus from other mussids is the asexual reproduction by extratentacular budding (Wells 1968). Polyps of a single colony are usually genetically identical with a high probability of having the exact structure and potential for physiological functions (Meester & Bak 1995) as the colony is derived from the budding of a single polyp (Jackson & Coates 1986). This is reflected in intratentacular budding, where parent polyps produce mature polyps of the same developmental stage (Sakai 1998). However, extratentacular budding by Blastomussa members produces developmentally ‘young’, small polyps that are mostly not fertile, unlike the parent polyps (Sakai 1998). In this case, the age of the polyp is reflected by their size, and thus egg production in polyps that displayed extratentacular and intratentacullar budding were different, following their maturity (Sakai 1998). This might suggest a low egg production for members of Blastomussa as compared to other mussids who bud intratentacullarly, thus explaining their rare occurences.

Reef-building corals tend to reproduce sexually by broadcast spawning, where their mature gametes are released into the coelenteron, and then spawned as a sperm and egg bundle into the water column via the mouth which then separates to allow for external fertilization, and then for embryogenesis to occur (Figure 7) (Baird et al 2009; Ruppert et al 2009). The highly seasonal synchronous broadcast spawning occurs annually and have been recorded to be multispecific (Babcock et al 1986). Gametogenic cycles of broadcast spawners are synchronized to potentially reduce predation (Babcock et al 1986) and thus increases survival and dispersal of the larvae to adulthood. Another method of sexual reproduction utilised by fewer reef-building corals is brooding, where the corals release developed planulae that are already competent for settlement. In this case, the life cycle does not include a free-swimming larva, thus potentially enhancing survival and hatching in a suitable habitat where they were brooded by successful adults (Brusca et al 2016).

Feeding

The life cycle of corals that develop indirectly includes a free larval stage. In the case of the Blastomussa, the larvae are likely planktotrophic (Ruppert et al 2009). This means that the eggs of the larvae are isolecithal, containing little yolk, and thus the larvae must feed on plankton in the pelagic seas to survive. Key to a successful larva is to feed to excess, such that they have sufficient nutrients to fuel their settlement and metamorphosis into a juvenile (Brusca et al 2016).

Once the larvae have transitioned into an adult polyp, they are polytrophic and have three feeding mechanisms – tentacular feeding, mucous-net suspension feeding, and obtaining nutrients directly from their symbiotic photosynthetic zooxanthellae. Cnidarians typically capture prey with their nematocyst-covered tentacles, thus stunning them in the process, and is then brought to the mouth to be ingested (Brusca et al 2016). In mucous-net suspension feeding, mucous is secreted from the mouth in thin strands or sheets which collect organic particulate matter in their surroundings. Cilia then directs the mucus back into them mouth of the polyp, along with the trapped food (Brusca et al 2016). Symbiotic zooxanthellae are able to photosynthesise in the day and supply their coral host with organic compounds such as glucose, providing potentially up to 50 percent of their energy requirement (Ruppert et al 2009) (see 'Symbiosis' in the 'Ecology' section).

Anatomy and Physiology

Morphology

Scleractinians are known for their massive calcareous skeletons. This calcium carbonate skeleton is secreted by epidermal cells located on the bottom half of the column. The skeleton is generally considered external as the coral sits on a non-living calcareous framework, although some biologist considers it an internal skeleton due to the thin secreting epidermis layer that covers the skeleton (Brusca et al 2016).

A corallite is the skeleton of a single polyp (Figure 8), which features the theca (outer wall), basal plate (floor), collumella (supportive skeletal process), and numerous septa which project inwards as support for the mesenteries of the polyp, which occupies the top surface of the corallite. As polyps grow, skeletal thickness increases (Brusca et al 2016).


 

Figure 8. Illustration of morphological features of a corallite of a solitary scleractinian coral. Adapted from Brusca et al (2016). 



B wellsi. polyps have 4 cycles of septa, with the first three of approximately equal size that meets the columella in the centre of the corallite, while the fourth is incomplete or heavily reduced (Figure 9) (Benzoni et al 2014). The septa of B. wellsi is made of multiple fan systems and possess finely granulated and dentated, lobed margins. The inner region of these margins of the septa and papillae forms the columella via trabecular processes. The lamellar structure in the centre of the corallite results in a bilateral symmetry of the columella, derived from the fusion of papillae (Figure 10) (Benzoni et al 2014). The corallite wall, bearing costae, is the site for the formation of epitheca, located just a few millimetres below its margin (Benzoni et al 2014).


  

Figure 9. (left) Side view of a B. wellsi corallite, with the septa cycle numbered. Image source: Benzoni et al 2014.
Figure 10. (right) Bilaterally symmetrical columella of a B. wellsi corallite. Image source: Benzoni et al 2014. 



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Figure 8
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Figure 9
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Figure 10

Nutrient Transport

Digestion in cnidarians occur extracellularly in the coelenteron, also known as the gastrovascular cavity, following the immediate consumption of prey (see ‘Feeding’ in the ‘Life History & Behaviour’ section) with the aid from enzyme-producing cells from the gastrodermis (Brusca et al 2016). As cnidarians lack a true circulatory system, distribution of partially digested material occurs in the coelenteron, a blind gut. Carbohydrates, polypeptides and fats from the partially digested compound are then phagocytosed or pinocytosed by nutritive-muscular cells, with intracellular digestion then completing digestion (Brusca et al 2016).

 

Figure 11. The general anatomy of a typical anthozoan, illustrating the coelenteron. Adapted from Brusca et al (2016). 



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Figure 11

Respiration & Excretion

The coelenteron is responsible for circulation in the polyp, where undigested waste products are expelled via the mouth of the polyp. Cnidarians are fairly structurally simple animals and lack specialised gas exchange and excretion organs. As the body walls of polyps are fairly thin, gas exchange from the internal and external body surfaces are possible as diffusion distances are short (Brusca et al 2016). Some metabolic waste is absorbed by the gastrodermis, which is then utilised by their symbiotic zooxanthellae (see ‘Symbiosis’ in the ‘Ecology’ section), and the rest, in the form of ammonia, simply diffuses through the body wall into the surrounding waters in the same way that gas exchange occurs (Brusca et al 2016).

Muscular & Nervous Systems

Most cnidarians have a non-centralised, diffuse nervous system. Within the animal kingdom, their neurosensory cells are mostly nonpolar and naked and thus considered the most primitive (Brusca et al 2016). They possess motor neurons and sensory neurons, with interneurons that connect the two together, and also the nerve nets at the base of the epidermis and the gastrodermis together (Brusca et al 2016).

Cnidarians possess myoepithelial cells, which form circular and longitudinal fibrils. Circular fibrils are made of epitheliomuscular cells and are associated with the gastrodermis, while longitudinal fibrils are associated with the epidermis. As most polyps of scleractinia are sedentary or sessile, they move mainly to capture food, using the epidermal muscles located in their oral disc and tentacles, or to withdraw their body away from an external stimulant, using the gastrodermal muscles in the column to contract their body (Brusca et al 2016). For polyps to ‘expand’ their tentacles and body, the gastrodermal muscles work in conjunction with the hydrostatic skeleton.

Cnidae

A unique defining feature to cnidarians is the presence of cnidocytes. These stinging cells are produced in cnidoblasts and develop from cnida in the epidermis or gastrodermis of the polyp to fully formed cnidocytes. Its many functions include locomotion, prey capture and immobilisation, and defence. Cnidocytes are present in the epidermis of all cnidarians and are in higher densities in tentacles. In some groups, they are present in regions of the gastrodermis (Brusca et al 2016).

Fluorescence

One of the most fluorescent animals in the oceans are corals. Some species have the ability to express a range of fluorescent proteins (FPs) and nonfluorescent chromoproteins (CPs) of vivid colours (Oswald et al 2007). In scleractinians, colour polymorphism results from the different levels of environmentally regulated expression of these FPs and CPs (Oswald et al 2007; Gittins et al 2015). The most well-known fluorescent protein is likely the green fluorescent protein (GFP), which is used in molecular markers, biosensors, fusion tags, and transcriptional reporters in molecular and cellular biology (Ben-Zvi et al 2015). The pigment produces a chromophore that absorbs short wavelength light and re-emitting it at a longer wavelength, producing the green fluorescence (Ben-Zvi et al 2015).

Corals and their zooxanthellae are more likely to experience more stress in shallower areas of reefs due to intense and excessive light exposure. One of the functions of GFP in shallow-water anthozoans is the photoprotection for their symbiotic zooxanthellae by re-emission or reflectance of photons from the algal pigments (Schlichter et al 1994; Gittins et al 2015). It was found that light intensity regulated the expression of GFP-like proteins in shallow-water anthozoans, with heat negatively affecting expression (D’Angelo et al 2012). This genetic framework thus allows corals to adopt either an expensive high-level of pigmentation, which would benefit them under light stress, or a low-level pigmentation, which would conserve resources for other uses (Gittins et al 2015).

Another function of these pigments is the facilitation of photosynthesis (Schlichter et al 1994). This occurs as the pigments enable the absorption of wavelengths of light that are usually poorly harvested by zooxanthellae for photosynthesis, also while enhancing the light supply to the zooxanthellae by the scattering and reflection of photons on the chromatophore pigment granules (Schlichter et al 1994). Other proposed functions of these pigments are the attraction of zooxanthellae (Hollingsworth et al 2004) and antioxidant function (Bou Abdallah et al 2006).



Mini Project

The B. wellsi specimens of this study were obtained from a specialist marine aquarium shop located in Wynnum, Queensland. The specimens were collected from the northern Great Barrier Reef, off Cairns. They were cut and cleaned to remove pests such as vermetid gastropods (Shima et al 2010) and pathogens associated with coral diseases like black band disease (Frias-Lopez et al 2004), then glued onto porcelain fragment tiles. The animals were then acclimatised to the aquarium tank before being purchased. They were then acclimatised to the holding tanks at the University’s aquarium at the Goddard Building before being examined.

Using a fluorescence microscope, I examined a B. wellsi polyp to determine if the species fluoresced naturally. I found that B. wellsi naturally fluoresced under two different wavelengths. Two filters were used, a fluorescein-5-isothiocyanate (FITC) filter with a narrow bandpass window in the blue spectral region (490-505 nanometers), and a Texas Red (Tx RED) filter with a narrow bandpass window in the green spectral region (560-580 nanometers).

Under the FITC filter, the polyp produced a green fluorescence at the mouth and at the mantle vesicles, mostly in their centres (Figure 12a). There were also some specks of green fluorescence observed on the oral disc of the polyp (Figure 12a). Under the Tx RED filter, the polyp produced a red fluorescence at the theca (Figure 13 b) and the mantle vesicles, mostly at their edges (Figure 12b). Unlike the green fluorescence produced, there were no traces of red fluorescence on the oral disc of the polyp (Figure 12a, b). Using Adobe Photoshop, I layered the images of the green and red fluorescence to obtain a complete fluorescence image of the polyp (Figure 12c & 13c).



 

Figure 12. Images of a B. wellsi polyp section showing the mouth, oral disc, and mantle vesicles, viewed under (a) a FITC filter, producing a green fluorescence, (b) a Tx RED filter, producing a red fluorescence, and (c) the combined images of (a) and (b), with the yellow indicating an overlap of the green and red fluorescence. 



 

Figure 13. Images of a retracted B. wellsi polyp, viewed under (a) a FITC filter, producing a green fluorescence, (b) a Tx RED filter, producing a red fluorescence, and (c) the combined images of (a) and (b), with the yellow indicating an overlap of the green and red fluorescence. 



As the two B. wellsi polyps obtained were of different colour morphs (Figure 1), I was initially interested to explore if they naturally fluoresced similarly or differently. However, the tissues of one of the polyps could not be obtained for viewing under the fluorescence microscope due to its quick disintegration into the water after being cut, and thus no comparison could be made. The observations of this project of the natural fluorescence of a single B. wellsi polyp may still possibly prove useful for future research, as there is scarce knowledge of this species apart from its morphology.



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Figure 12
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Figure 13

Biogeographic Distribution

B. wellsi has been observed and recorded from the Red Sea, the Indian Ocean, and the western Pacific Ocean (Benzoni et al 2014). A map of the species’ native range is illustrated (Figure 14), along with a map of suitable habitats for the species (Figure 15).




Figure 14.
Native range of Blastomussa wellsi, based on data points of recorded sightings from the Ocean Biogeographic Information System. Image source: www.sealifebase.org/summary/Blastomussa-wellsi.html






Figure 15.
Map of suitable habitats of Blastomussa wellsi, based on environmental conditions of the oceans and the species’ known environmental requirements. Image source: www.sealifebase.org/summary/Blastomussa-wellsi.html


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Figure 14
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Figure 15

Evolution and Systematics

The approximate 1400 known species of scleractinians (true stony corals) are placed among 27 families from classical morphological taxonomy (Kitahara et al 2010). Phylogenetic studies of marine speciation are more frequently using corals as a model (Vollmer & Palumbi 2002). Despite the use of molecular phylogenetic methods, however, most aspects of coral evolution still remain unresolved. Macroscopic skeletal characteristics are the primary basis of traditional scleractinian coral systematics at genus level (Veron & Pitchon 1980), but the recent discovery of the considerable variance between this method and molecular phylogenetic studies has subjected the former to intense scrutiny (Romano & Palumbi 1996). The increasing interest in reef-building coral evolution and systematics is driven by the concern for their conservation (Kerr 2005). In Kerr’s (2005) study to create a supertree of scleractinia, he found that a third of phylogenetic effort within this order of anthozoans only focussed on the most diverse genera Acroporidae

Although the Blastomussa genus was initially classified into the mussidae family Ortmann, 1980, a revision of this family was made by Budd et al (2012) following molecular analyses and microstructural research. Mussidae now only occurs in the Atlantic Ocean, thus placing Blastomussa in the Lobophylliidae family Dai & Horng, 2009 (Benzoni et al 2014). However, their distant relation to the rest of the Lobophylliidae family eventually led to their classification into the Plesiastreidae (Dai & Horng 2009) or incertae sedis (Budd et al 2012), which means ‘of uncertain placement’ due to the undefined broader relationships between other taxa.

Kingdom – Animalia
Phylum – Cnidaria
Class – Anthozoa
Subclass – Hexacorallia
Order – Scleractinia (true stony corals)
Family – Plesiastreidae / incertae sedis
Genus – Blastomussa
Species – B. Wellsi

From Kerr’s (2005) supertree, Blastomussa is positioned in the clade Faviina + Meandriina (Figure 16), one of the three clades further derived from the basal clade Robusta of Romano and Palumbi (1996). This clade comprises mostly of species from faviidae, including mussidae. Although the exact position of the Blastomussa genus is not well resolved, the monophyly of five Blastomussa species was shown in a study by Benzoni et al (2014) using rDNA and mitochondrial gene COI phylogeny (Figure 17). The most closely related species to B. wellsi was B. vivida, both possessing larger corallites, fleshier polyps, and higher numbers of strongly dentated septa as compared to the other three Blastomussa species (Benzoni et al 2014).


 

Figure 16. Summary of the phylogenetic supertree of scleractinia, adapted from Kerr (2005). The phylogenetic position of Blastomussa genus is highlighted in red. 



 

Figure 17. Phylogenetic tree of mitochondrial gene COI reconstructed with Bayesian inference, adapted from Benzoni et al (2014), with the red highlighted section indicating the evolutionary position of Blastomussa wellsi. 


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Figure 16
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Figure 17

Conservation and Threats

The status of Blastomussa wellsi is Near Threatened on the IUCN Red List (2008). The biggest threat to B. wellsi is the commercial harvesting for the aquarium trade, with Indonesia leading in export of an annual quota of 3,800 live pieces in 2005 (IUCN Red List, 2008).

Another major threat to corals is climate change. Since the industrial revolution, approximately 142 billion tonnes of atmospheric carbon dioxide, a third of all emissions (Feely et al 2009), has been absorbed by the oceans (Simpson et al 2011). This has led to ocean warming, ocean acidification and increased intensity and frequency of storms, which has led to secondary effects in corals such as increased vulnerability to disease, inhibition of proper skeleton growth, and mass bleaching events. With the warming of the ocean, the mean temperature of the ocean is predicted to rise by 12°C by year 2100 (Scott 2016). Ocean warming has already caused mass bleaching events (ARC Centre of Excellence for Coral Reef Studies, 2017) as the increase in temperature by 1°C causes corals to lose their symbiotic algae (see ‘Symbiosis’ in the ‘Ecology’ section), resulting in bleaching that may potentially lead to death of the coral (Doney et al 2012). The increase in carbon dioxide levels has led to ocean acidification, where ocean pH, carbonate ion concentration, and aragonite saturation state has all declined (Orr et al 2005), resulting in the severe inhibition of proper skeleton growth in corals (Doney et al 2009). More frequent and intense storms are predicted to occur with climate change, which has the capacity to severely damage coral reefs; Between 1985 and 2012, the Great Barrier Reef lost 42% of its corals due to storms (De’ath et al 2012).

Another large threat to coral reefs is coral disease (IUCN Red List 2008). The increase in vulnerability of corals to disease due to climate change, together with the increase in number of coral species affected by diseases and types of diseases, increases the risk of coral death (Green & Bruckner 2000). Bruno et al (2007) showed a correlation between increased ocean temperatures and increased coral disease levels on the Great Barrier Reef.

A map of the possible year 2100 native range of B. wellsi based on the IPCC A2 emission scenario (IPCC, 2000) is illustrated below (Figure 18). The decrease in red area in comparison to its current native range (Figure 14) reflects a large decline in species range. Although the combined impact of these threats specifically to B. wellsi is unknown, coral reefs are undeniably at a high risk of collapse. There are currently no known conservation efforts for B. wellsi.


 
Figure 18. Possible year 2100 native range of Blastomussa wellsi, modelled based on IPCC A2 emissions scenario. Image source: www.sealifebase.org/summary/Blastomussa-wellsi.html
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Figure 18

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