A Synthesis Paper on Seahorse (​​Hippocampus sp.

)​ Reproduction and Mating Behaviour

By

Ma. Victoria Ty Cruz

2015-89009

To

Asst. Prof. Patrick C. Cabaitan, Ph.D.

University of the Philippines, Diliman

December 6, 2018
 ABSTRACT

The process of reproduction and parental investments of both time and energy vary

widely among species (Stoelting, 2010). In nature, female investment and maternal care in

offspring is the norm and is also much more salient than male investment and paternal care.

When it comes to bending gender stereotypes, seahorses ​(Hippocampus sp.) would have to be

one of the most extreme examples. Male seahorses take on a role that is extremely uncommon in

the animal kingdom and nature in general; they get pregnant and deliver their offspring. In line

with that, the three main objectives of this study are to: 1) describe the reproductive process of

seahorses, 2) explain the structures involved in reproduction, and 3) understand mating

behaviour in seahorses. The paper therefore synthesizes existing literature to get a critical

overview on seahorse reproduction. From the literature, it was found that both male and female

seahorses possess specialized and unique structures, the brood pouch and the ovipositor,

respectively. The brood pouch is an appendage found near the male’s tail, which provides

aeration, protection, osmoregulation and nutrition to the embryo or offspring after the females

deposits eggs during mating (Lin et al., 2012). It was also found that mating behaviour of

seahorses consist of courtship phases and distinct behavioural changes during the day of

copulation (Lewis & Masonjones, 1996). Lastly, even though there is a high degree of paternal

care, courtship roles are not necessarily reversed in seahorses. Males are still more active in the

courtship process and compete more intensely than females for partners (Jones, 2004).

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 INTRODUCTION

Seahorses are bony fishes (teleosts), complete with gills, fins and a swim-bladder (Lourie

et al., 1999). They are characterized by a sparse distribution, low mobility, small home ranges,

low fecundity, lengthy parental care and mate fidelity (Foster & Vincent, 2004). Seahorses are

usually found in warm shallow, temperature and tropical waters. What is very unique and notable

about seahorses is that they are known to be the one of the only species where is it is the male

that become pregnant.

Aim of Study

This paper aims to first provide a brief overview on seahorse (​Hippocampus sp.​) life

history and ecology. More importantly, the main focus of this paper is on seahorse reproduction

and male pregnancy. Male seahorses do not follow traditional roles in reproduction, as they are

the ones that carry and deliver the offspring. It is the male that provides all postfertilization

parental care, as the male seahorse has certain morphological and physiological adaptations to

nourish the developing embryos. Therefore, this paper aims to synthesise existing literature on

seahorses, and thus the three main objectives of this study are the following: 1) to describe the

reproductive process of seahorses, 2) to explain the mechanisms and structures involved in

reproduction, and 3) to understand mating behaviour and sex roles in seahorses.

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 SEAHORSE TAXONOMY

A seahorses is a type of fish that belong to the genus ​Hippocampus o​ f the family

Sygnathidae. Today, there are 54 extant species of seahorses that are taxonomically recognized

as valid, and all within the single genus ​Hippocampus (Cohen et al., 2017). The entire family

Sygnathidae falls under order Gasterosteiformes, which consists of 55 genera of pipefishes,

pipehorses, and sea dragons (Kuiter, 2000 as cited in Foster & Vincent, 2004). Syngnathid fishes

are a group of 232 species, which exhibit a wide diversity of brooding types and structures

varying in their complexity and location on the male body (Wilson et al. 2001; Kuiter 2000, as

cited in Stoelting, 2010).

The family Syngnathidae (seahorses and pipefishes) is the sole vertebrate group where

the embryonic development occurs within a special pouch in males (Herald, 1959, as cited in Lin

et al., 2012). Fishes in this family Syngnathidae show a wide variation in the degree of male

anatomical and physiological specialization for paternal care, which is evidenced by the different

morphological structures that these fishes possess, (Herald, 1959; Vincent et al., 1992 as cited in

Lewis & Masonjones, 1996). Amongst the fishes in the family, seahorses exhibit the greatest

specialization of paternal care, because it is the female that transfers eggs directly into an

abdominal brood pouch on the male, where fertilization takes place (Fiedler, 1954, as cited in

Lewis & Masonjones, 1996).

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 SEAHORSE MORPHOLOGY

All seahorses are bony fishes that have gills and fins; however, unlike most fish, they do

not have scales. They consist of a horse-like head (hence, their common name “Seahorse”),

which is positioned at a right angle to their body. Seahorses also have a crown, known as a

coronet, which is so unique that it gives rise to a seahorse’s identity. Seahorses do not have teeth;

instead, they have a long tube-shaped snouth, which has a powerful suction so they can swallow

and suck their food whole. They also have a prehensile tail, which is used to grab vegetation in

the sea floor. They wrap their tail around the stems of seagrass so that they can hang onto

substrate, thereby helping them to avoid being swept away by strong currents. Their eyes also

move and swivel independently from one another, which helps them in looking out for potential

predators. Their skin is stretched over rectangular bony plates, which are visible as rings around

their body, especially around the trunk and tail (Lourie et al, 2004).

Juveniles tend to have larger heads relative to their bodies, compared to adults. They also

are slimmer and spinier in form, and their coronets are relatively higher. Additionally, adult

seahorses do not have pelvic and caudal fins, but they retain two small ear-like pectoral fins,

which are used for stabilization and steering, (Foster & Vincent, 2004). The dorsal fin, which is

located on the lower part of the body, is a small appendage which helps propagate the seahorse’s

movement forward. The pectoral fins, which are located on the head near the gills, controls the

side to side movement and therefore enables the seahorse to turn from one orientation to another.

Lastly, in terms of size of a seahorse, their full length can range from a quarter of an inch to a

foot or more depending on the species.

4
 ​ igure extracted from Loh et al., (2014) as cited
Figure 2. The parts of a seahorse, ​H. barbouri. F

in Foster & Apale (2016).

As seen in Figure 2, the major difference between male seahorses and female seahorses is

that, in females, the belly does not extend past the bottom of the dorsal fin. Most importantly,

males possess a brood pouch, located near the male’s tail. This is a pouch wherein fertilisation

takes place and where the offspring are carried by the male. On the other hand, females possess

an ovipositor, a protrusion found at the bottom of the torso, which is used to insert and deposit

unfertilised eggs into the male’s brood pouch.

Males also generally have longer tails, while females have longer trunks. Moreover,

seahorse weights vary; however, they increase when females are carrying eggs or when males

are carrying their offspring (Louirie et al, 2004). Moreover, female seahorses usually have a

more vibrant colouring compared to male seahorses, which have a more subdued coloring,

because this helps females attract a mate.

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 SEAHORSE ECOLOGY

Habitat

Most seahorses live in temperate and tropical regions. They are found in relatively

shallow parts of the water, usually seaweed beds, since the water is warmer. Seahorses prefer

environments with still waters or waters with a very slow current, because seahorses are weak

swimmers. Because of this disadvantage, they stay in shallow waters where their prehensile tail

can easily grasp onto rocks or grass for when they need to rest. Thus, they are usually found in

soft-bottom areas that also serve as habitats to corals, sea sponges, seaweed, and other grasses.

Many species in temperate and tropical regions live among seagrasses or eelgrasses while others

occur in flooded mangrove forest (Foster & Vincent, 2004). The local environment in these

shallow waters ranges from marine to estuarine habitats, including coral reefs, mangroves, and

temperate seagrass beds (Kuiter, 2000, as cited in Lee, 2013).

Camouflage Ability

Seahorses posses the excellent ability to change color, which helps facilitate both prey

capture and predator avoidance (Foster & Vincent, 2004). When they undergo various color

changes, seahorses stay immobile for a long time, and they spend a few days or weeks to match

their environment and surroundings better. They can also undergo short-term color changes,

which usually occurs during courtship and other intra-species interactions. This ability to change

color also helps the seahorse in hiding and protecting themselves from predators. Seahorses

usually grow long skin appendages so that they can blend in better with the algae around them.

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However, due to their bony armour, seahorses do not look as appetizing, and therefore do not

have as many predators as other marine animals.

Feeding

Seahorses very much depend on their vision in their search for prey. Because their eyes

swivel independently of one another, seahorses have the ability to look into different directions

as they swim around. Though relatively small creatures, seahorses actually have very large

appetites. Their diet consists of plankton, but it is dominated by small crustaceans, such as

copepods, amphipods, caridean, and mysid shrimp. Because they ingest their prey through a

tubular snout and because they lack teeth, seahorses eat their food whole and intact (Woods,

2002). Moreover, this also means that seahorses can only eat small quantities of food at a time,

and therefore they feed continuously to ensure that they get enough food and nutrients. When

feeding within the water column, they wait until prey comes close to their mouth. When this

happens, the prey are drawn up into the long snout with a rapid intake of water (Foster &

Vincent, 2004). Seahorses are also known to be ambush predators, as they lie around waiting for

prey. They heavily rely on stealth and camouflage to approach within striking range of their

prey, or allow their prey to move within striking range (Flynn & Ritz, 1999, as cited in Woods,

2002).

Most seahorses are diurnal with the majority of their feeding behaviour occurring during

daylight hours (Kuiter, 2000, as cited in Lee, 2013). Due to their prehensile tail, they wrap

themselves around stationary objects (such as seaweeds). Because their feeding is visually

guided, seahorses use their eyes to orient them towards any possible prey (Lee, 2013).

7
 SEAHORSE REPRODUCTION

Mating Behaviour

The mating behaviour and rituals of male and female seahorses is quite fascinating. As

seen in Figure 3, the courtship ritual begins when the male and the female take part in seductive

dances together. This consists of the seahorses intertwining their tails with one another and

swimming together. They mirror each other’s movements and can also can change color as they

move together. Both the male and female will engage in this prolonged and elaborate courtship

ritual, which can last several hours (Jones, 2004).

Once the courting stage is over and if the conditions permit, the male will ready his

pouch by pumping water into it. The couple will stop their dancing and halt in the middle of an

open water column. Their bodies will be held close together, with the horse-like heads forming

the shape of a heart, and their snouts and bellies will touch one another. It will take several tries

for them to get into the right position, since the female has to be just above the male.

Once the male and female finally get into the perfect position, the female will deposit the

unfertilised eggs using the ovipositor (an appendage that protrudes out of the bottom of the

female’s torso) into the male’s brood pouch. This can be seen in Figure 4. The walls of the brood

pouch provide maximum surface area so that every embryo can embed in the soft tissue. The

actual transfer of eggs from the female’s ovipositor into the male’s brood pouch only takes a few

seconds.

When the eggs have been transferred into the brood pouch of the male, intra-pouch

fertilization occurs, which is when the male releases his sperm inside the pouch to fertilise the

8
eggs. Because sperm cells are shed above the brood pouch and enter the pouch lumen, sperm-egg

union is much more efficient as it occurs without the necessity of extended sperm movement and

lengthy zygotic migrations to the site of implantation, (Watanabe et al, 2000 as cited in Stoelting,

2010).

At this point, the female’s investment in the offspring is complete, but the male is left

with a pouch full of growing offspring. Unfortunately, not all the eggs deposited in the male’s

pouch successfully complete the development and hatch as juveniles (offspring).

Since seahorses have a “male-bears-the-young” reproductive strategy, the male carries

the developing embryos for a period of several weeks. As the pregnancy lengthens, the pouch

becomes more and more of a serious burden for the male. It bulges and becomes heavier,

therefore males do not move so much while pregnant.

Even though the male is solely responsible to take care of the developing embryos, the

female, although she provides no parental care, visits the male. The female checks on the

progress of the male’s pregnancy with a brief visit each morning. She does this so she can verify

on the status of the pregnancy and also know when to have a new set of eggs ready again, so that

she can re-impregnate him once he has given birth to their offspring.

Once released from the brood pouch, the newly-born offspring are completely on their

own. Because there is no care from either parents, seahorses have a low survival rate. They are

easily carried away by currents, as they cannot latch onto substrate with their prehensile tail.

Depending on the species, male seahorses can deliver up to 1552 juveniles (Teixeira and Musick,

2001 as cited in Lin et al., 2012). Lastly, within the same day of giving birth, the male can get

ready to start his next pregnancy (Jones, 2004).

9
 Figure 3. Seahorses ‘flirting’. Photo by Norbert Wu, extracted from The Guardian.

Figure 4. Seahorses mating. Photo by George Grall, extracted from The Guardian.

Phases of Courtship

Lewis & Masonjones (1996) conducted a study on the courtship behaviour in the dwarf

seahorse, ​H. zosterae.​ During courtship interactions, seahorses were positioned side by side

facing in the same direction, and both males and females exhibited a characteristic posture,

which consisted of an erect body with their head inclined downwards.

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 More importantly, it was found that there were four distinct phases of courtship, which

are marked by distinct behavioural changes and differences in the intensity of courtship. The first

phase is the “initial courtship” phase, which occurs for one or two mornings before the day of

copulation. This is characterized by reciprocal quivering, which consists of rapid side-to-side

body vibrations displayed alternately by males and females. This courtship occurred for either

one or two days, preceding the day of copulation; however, it was observed that courtship

duration was significantly longer on the first day. The first phase is the only phase that takes

places preceding the day of copulation; as the rest of the phases are restricted to the day of

copulation.

The second phase is known as the “pointing-pumping” phase, which takes place on the

day of copulation. This phase is distinguished by females exhibiting a certain pointing posture,

during which the head of the seahorse is raised upward. When the female goes into the pointing

posture, she learns her body toward the male, who simultaneously leans away.

The third phase is known as the “pointing-pointing phase,” because this is wherein the

males assumes the pointing posture in response to female pointing. It is also during this phase

wherein the males and females initiate courtship by approaching each other equally often, and

they tended to brighten simultaneously.

The fourth and final phase of courtship, known as the “rising and copulation” phase,

includes a series of five to eight courtship bouts. Bouts during this phase were distinguished by

rising, since seahorse pairs rose upward together in the water column. Once they rise in the water

column, the female transfers her yolk-rich eggs directly into the male brood pouch during a brief

midwater copulation.

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 In successfully mating pairs, males initiated courtship by approaching the female and

brightening first. In the two pairs that failed to mate (from the 19 seahorse pairs that were

observed), females made the initial approach and brightened first, and the male responded by

brightening and then quivering. Lastly, it was observed that courtship activity levels increased

from relatively low levels during the first courtship phase to highly active courtship on the day of

copulation (Lewis & Masonjones, 1996).

Male Brood Pouch

The brood pouch is a brooding structure that can be found in male seahorses (as seen in

Figure 5), which is analogous to the womb of a female mammal. During pregnancy in mammals,

the embryo is implanted in the uterus of the mother, and a physiological connection between the

mother and offspring is made possible by the placenta and umbilical cord. Similarly, in

seahorses, there are paternal tissues that surround the eggs within the male’s brood pouch,

therefore allowing blood flow through this tissue and thus providing a placenta-like connection

between the male and his offspring (Jones, 2004).

This complex organ acts like a chamber, as it can alter its composition to resemble the

characteristics of the surrounding ocean water. The male controls the concentrations of ions and

oxygen within the pouch and the male also provides the obvious protection afforded by the outer

pouch wall (Jones, 2004). The pouch can regulate factors such as blood flow, water salinity, and

temperature. In fact, it was found that the osmolality of pouch fluid changes substantially during

incubation, as it increases from that of paternal blood at the time of fertilization, to that of the

surrounding marine environment later in development (Leiner 1934, Linton and Soloff, 1934, as

12
cited in Stoelting, 2010). Also, it was found that potential reproductive rate of seahorse ​H.

erectus ​increased with the brood pouch development (Lin et al., 2012).

While pregnant, the male seahorse monitors the the salinity within the pouch to match the

salinity of the environment, so that the offspring can already be accustomed to such conditions

while they are stilled inside his pouch. Therefore, the male seahorse slowly allows seawater into

his pouch so that the embryos will become used to such conditions before having to swim all

alone in the environment once they are born. Moreover, the male brood pouch epithelium is

highly vascularized and provides gas exchange and osmoregulation for developing embryos

throughout gestation (Boisseau, 1967; Linton and Soloff, 1964 as cited in Lewis & Masonjones,

1996).

Figure 5. The brood pouch in full display in two pregnant male seahorses. Photo by Clive

Bromhall, extracted from The Guardian.

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Reproductive Rate and Offspring Survivorship

In terms of the dynamics of the reproductive rate of seahorses, there are various factors

that affect seahorses’ potential reproductive rate and survivorship of offspring. One study found

that in the seahorse ​H. kuda,​ parental size and age could affect the number, survivorship and the

growth of the offspring, (Dzyuba et al., 2006, as cited in Lin et al., 2012). Moreover, factors such

as mate competition and physical interference during courtship and egg transfer also have to be

considered when estimating the offspring number from parents.

Another study by Lin et al. (2012) found that young pairs of seahorses ​H. erectus had

smaller brood size, poor offspring survivorship and growth, when compared with those from

older pairs. Reproductive efficiency was also found to have improved over successive births by

the males, since the mean brood size increased significantly from the 1st birth to the 6th birth.

Moreover, pairs of ​H. erectus ​had short greeting and re-mating durations before or after each

copulation, which significantly improved the reproductive rate. Lastly, this study also found that

the survivorship of offspring released older seahorses (i.e. 2 year old males) was higher than that

from younger seahorses (i.e. 1-year old males), which is probably due to the increased size of

brood pouches of the old males, or better function of the pouches in old males (Lin et al., 2012).

Mating Patterns

Seahorses are monogamous, which means they only mate with one partner. Monogamous

relationships are actually rare for fish, and thus this is another trait that makes the seahorses

unique. Seahorses are not only monogamous within the course of a single breeding episode, but

14
they maintain pair bonds that persist across mating events within a breeding season, therefore

constantly returning to the same partner, (Kvarnemo et al, 2000, as cited in Jones, 2004).

Reproductive potential of both sexes is positively correlated with body size. Seahorse

mating involves assortative pairing by size, which means that large males mate with large

females, while small males mate with small females. Seahorses benefit from such an

arrangement since larger males have larger brood pouches, while larger females produce more

eggs. If a large male mates with a small female, the female will not produce enough eggs to

completely fill the male’s brood pouch; and vice versa, if a small male mates with a large female,

the male’s brood pouch will be unable to accommodate all the eggs produced by the female. In

both cases, either the male or the female in the pair is wasting some reproductive potential

(Jones, 2004); therefore such an arrangement fully maximizes the reproductive potential of both

individuals in the relationship.

This is very much in line with the study by Lin et al. (2012), who found positive linear

correlations between male pouch size and the number of offspring produced, and as well as

female standard body length and the number of offspring produced. Generally, the body size of

offspring was correlated with the brood size, gestation time and nutrient supply in the same

males and females’ investment (Lin et al., 2012). Lastly, it was also found that larger males do

not necessarily release larger-sized offspring than those from smaller males.

Evolutionary Explanations

It is imperative to look into the evolutionary explanation of the reproductive formation in

seahorses, because their form of reproduction is truly specialized and unique in the animal

15
kingdom. There is still no clear-cut reason to explain why seahorses have evolved this way, but

scientists postulate male seahorses take charge of the pregnancy because it gives them full

reassurance that all the babies the male is ​caring for, are in fact, definitely his own. Because the

eggs are first deposited from a female’s ovipositor into the male’s brood pouch, it is only when

these unfertilised eggs are in the male’s brood pouch, that the male will release his sperm into the

pouch to fertilise the eggs.

Additionally, scientists have postulated that seahorses have evolved into having the male

carry babies because this gives them the ability to create more babies quickly. Once the female

has deposited all her eggs to the male, it gives time for the female to make more eggs. The

female expends time and energy in making the eggs, especially since the female uses this energy

to fill the egg casings with nutrients, which will help the developing young mature. Therefore,

this arrangement wherein the male is the one that carries the developing brood is actually an

advantage because it enables for the pair to distribute the work needed in the reproduction

process. While the male is carrying one set of eggs, the female can already be working on

making a new set of eggs for the next pregnancy. This is more efficient since once the male gives

birth to the first set of eggs, the second set of eggs will be ready and can be implanted in the

male’s brood pouch soon after.

A study looking into the evolution of male pregnancy using genetics posits that complex

structures with skin or pouch-like folds covering attached eggs can be traced from the basal

lineage of Gastrophori, (Stoelting, 2010). This originally was a simple brooding structure,

wherein there was an open gluing area for eggs at the abdomen, but this has evolved into the

male brood pouch that is currently seen in male seahorses today. In the family Syngnathidae, the

16
different lineages are classified based on the location of the male brood pouch; on the tail

(Urophori) and on the abdomen (Gastrophori). In the Urophori, a set of morphologically more

complex pouch types evolved in a rapid diversification between 20 and 52 MYA (Patterson

1993; Teske et al. 2003, as cited in Stoelting, 2010). However, close similarities in terms of male

pregnancy within the two lineages suggest that there maybe similar genetic mechanisms that are

responsible for generating these structures.

Various researches have also tried to use the sexual selection theory to try and explain the

mating behaviours of seahorses. Sexual selection theory predicts that how much investment is

given by both males and females in their offspring is a primary determinant of sexual selection

intensity and patterns of courtship behavior (Jones, 2004). Traditional sex roles during courtship

consists of males playing a much more active role in initiating courtship, males competing for

access to females, and females being selective about mates (Trivers, 1972; Williams, 1975;

Gwynne, 1991, as cited in Lewis & Masonjones, 1996). Therefore, in species wherein male

investment exceeds that of females, the sexual selection theory predicts that these traditional

courtship patterns should be reversed. This is also supported by the parental investment theory,

which predicted that in species with high male investment in offspring, females might compete

for access to males (Andersson, 1994 as cited in Jones, 2004). If it is the male that provides most

parental care to the developing young, this becomes very costly and therefore they will be too

busy to care for offspring that they will not be available to mate.

Therefore, Lewis & Masonjones (1996) posit that because seahorses have a high degree

of morphological specialization for paternal care, seahorses can thus be used to test the

prediction that relative parental investment determines sex roles during courtship. However, they

17
found that seahorses do not appear to show a reversal of traditional roles, in spite of their

extreme paternal care. By observing the courtship phases in seahorses, it was found that the

males are the ones that more actively initiated courtship on the days preceding copulation, and

also males were found to compete more intensely than females for access to mates. This

therefore indicates that these seahorses are not courtship-role reversed, as has previously been

assumed by the sexual selection theory and parental investment theory.

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 CONCLUSION

Male pregnancy is an extreme and specialised form of paternal care which are very much

unique to syngnathid fishes. Seahorses are remarkable and beautiful organisms that are very

much a unique species in terms of how they produce and the mating behaviours involved in the

reproduction process. Seahorses are stereotype-breakers in nature, because unlike what is often

seen with many other species, fertilization takes place inside a male’s brood pouch and they also

carry and nourish the offspring.

Moreover, several studies have shown that even though seahorses break the norm in

terms of who invests more care in their young, courtship roles are not necessarily reversed in

species wherein there is a high degree of paternal care (Lewis & Masonjones, 1996; Jones,

2004). In nature, traditional courtship roles dictate that is it the female that carries and nourishes

the developing young, and thus males are more active in courtship and males compete for access

to females. However, even though the roles are reversed wherein it is the male that carries and

highly invests highly in their developing young, it is still the males that are more active in the

courtship process and even on the day of copulation in seahorses.

It is also important to note that seahorses have the most-complex pouch structure and also

undergo the most significant physiological changes during embryo incubation. Moreover, it is

evident that the role of the seahorse pouch is far more than just a big of protecting the developing

young, as there is evidence for osmoregulatory, aerative, nutritive and possible

immunoprotective roles of the male brood pouch (Stoelting, 2010).

However, seahorses are actually an understudied species in the realms of marine sciences,

as there are still major gaps in the literature on seahorses. In fact, reproduction has also only been

19
studied in a fairly small number of seahorse species (Jones, 2004). Our understanding of

syngnathid reproduction, especially when compared to mammalian pregnancy, is still very basic

by comparison. Therefore, researchers and scientists alike should further study the compelling

morphological structures (such as the brood pouch and ovipositor) and the underlying processes

of pregnancy in seahorses. Lastly, there is a lack of evolutionary explanations as to how and why

seahorses have evolved into a species wherein male investment exceeds that of females. Because

there are only few studies on the topic of mating competition and sex roles, additional research is

certainly warranted.

Word Count: ​4832

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