Gabriella Juliane L.

Maala
MS 260 Case Study

The potential of interspecific hybridization as a means to enhance

coral reef resilience and biodiversity

Introduction
Corals are important reef-building organisms that exhibit a symbiotic relationship with
microalgae Symbiodinium. Unfortunately, as atmospheric CO2 levels increase rapidly, oceans
acidify and consequently destroy calcareous structures like reef-building corals (Chan &
Connolly 2013). Alongside ocean acidification is increasing ocean temperature due to global
warming which causes corals to expel their associated microalgae – a phenomenon called coral
bleaching (Hoegh-Guldberg 1999). According to several researchers (Eakin et al. 2016; Heron
et al., 2016; Hughes et al., 2017, and 2018 as cited by Chan et al. 2018), multiple mass
bleaching events have occurred for the last three decades, and unfortunately, it is expected to
have at least one mass bleaching event yearly before the century ends (van Hooidonk et al.
2016 as cited by Chan 2018).

Traditional approaches to conservation like habitat restoration and establishment of marine

protected areas have been effective to some extent (e.g., Selig and Bruno 2010, Sala and
Giakoumi 2018, Giakoumi et al. 2018). However, these efforts may not be enough to ensure
the ability of organisms to withstand rapidly aggravating stressors and anthropogenic
disturbances (Seastedt et al. 2008, Hobbs 2013, Hughes et al. 2017). To increase the likelihood
of success, it is thereby imperative that modern restoration strategies take into consideration
relevant information like current ecological characteristics of species, conditions of future
available habitats, and adaptive potential of the species (Becker et al. 2013). Alternative
approaches have consequently been proposed as ways to accelerate both the recovery of
depleted populations and the adaptation of populations to fast changing habitats (van Oppen et
al. 2015), one of which is assisted evolution. Assisted evolution mainly aims to enhance
resilience-related traits and hasten evolutionary progress of organisms by altering the genetic
makeup of organisms (van Oppen et al. 2015). Several approaches to facilitate assisted
evolution of reef-building corals include (i) preconditioning by exposure to nonlethal stress,
(ii) altering associated microbial communities, (iii) selective breeding, and (iv) genetic
engineering of microalgal symbionts (van Oppen et al. 2015). Interspecific hybridization,
which could be categorized as selective breeding, will be the focus of this case study.

Interspecific hybridization occurs when two genetically distinct individuals mate, and their
gametes are compatible enough to overcome post-zygotic barriers and produce a viable hybrid
offspring. Hybridization is often used to increase genetic variation, promote evolvability, and
induce hybrid vigor, which is the enhancement of traits like adaptive potential that are
otherwise hindered by intraspecific gamete exchange (Hoffmann and Sgrò, 2011; Becker et al.,
2013; Carlson et al., 2014; van Oppen et al., 2015; Hamilton and Miller, 2016; and Meier et
al., 2017 as cited by Chan et al. 2018). Proposed theories on the molecular mechanisms of
hybrid vigor include: (1) dominant alleles masking recessive alleles (Davenport 1908), (2)
novel allelic combinations inducing better heterozygote traits (Shull 1908), (3) unknown
interactions among newly combined loci (Crow and Kimura 1970), and/or (4) unique
interactions between and among different levels of gene control (Kaeppler 2012), thereby
influencing the fitness of hybrids.

Significance
Interspecific hybridization has always been considered a controversial approach to
conservation, and rightfully so given the possible dangers it can cause to existing wildlife such
as (but not limited to) loss of genetic biodiversity and outbreeding depression (Laikre et al.
2010). Furthermore, there is currently very limited studies on coral hybridization using this
method for conservation. However, with the limited studies we have, successful hybridization
and fitter hybrid offspring have been reported several times as will be discussed later. These
findings imply the potential of interspecific hybridization to aid in enhancing reef resilience
and biodiversity. As such, this case study explores the potential of interspecific hybridization
as a reef conservation approach and whether studies on it are worth pursuing in the future.

Objectives
This case study aims to explore the potential of interspecific hybridization as a means to
enhance coral reef resilience and biodiversity. It specifically aims, first, to present (1)
arguments both for and against interspecific hybridization as a reef conservation approach, (2)
coral hybridization studies comparing traits of hybrids and purebred parents, and (3)
predictions on how interspecific hybridization might influence reef resilience and biodiversity.
Considering what has been read and personal insights consequentially formed, it then
specifically aims to determine whether hybridizing and transplanting reef-building corals is
worth exploring in the future. Finally, if the previous objective is in the affirmative, concrete
steps to advance coral hybridization research will be proposed.

Key concerns on the use of interspecific hybridization and assisted evolution for
conservation
Even if interspecific specific hybridization can increase resilience and biodiversity, it can also
cause the exact opposite. The main concerns of using hybridization for conservation is the
possibility of outbreeding depression, loss of current genetic diversity, and the ethics and
likelihood of success in doing so. Here, risks of interspecific hybridization will be presented,
as well as counterarguments to the frequency or likelihood that these risks will occur and/or be
overcome.

Outbreeding depression
A threat against potentially gaining innate resilience and stress tolerance of potential hybrids
is outbreeding depression. Hybridization can create allele combinations with deleterious
effects, thereby causing outbreeding depression (i.e., fitness of a hybrid offspring is
significantly reduced) (Hamilton & Miller 2016). However, outbreeding depression is also only
likely to happen if hybridization is between populations that are geographically distant or
between species with significantly varied life history or phenological traits (Hwang et al. 2012,
Whiteley et al. 2015 as cited by Chan et al. 2018). In the case that it has occurred, outbreeding
depression can be temporary and eventually removed through natural selection (Hamilton &
Miller 2016). Consequently, outbreeding depression is not as common as originally thought
(Frankham 2015, Hoffmann et al. 2015 as cited by Hamilton & Miller 2016), and that cases of
inbreeding depression are more frequently observed to occur (Chan et al. 2018).

Loss of current genetic diversity

Reduction in standing biodiversity is another risk that occurs when aliens such as introduced
hybrids are introduced to a community, and this can occur in several ways. For instance, a
hybrid with greater relative fitness or novel traits may gain a competitive advantage over the
natives (Muir & Howard as cited by van Oppen et al. 2015). Eventually, hybrids can promote
loss of current genetic diversity through either: (1) causing an invasion that depletes native
populations, or (2) accidentally mating with native populations and disrupting their established
fitness-related gene complexes (Laikre et al. 2010). Although these are all possible aftermaths
of interspecific hybridization, loss of biodiversity is said to be more probable if products of
rather extreme methods like genetic engineering are the organisms involved (van Oppen et al.
2015). Interspecific hybridization is still considered a relatively less drastic and widely
accepted method (van Oppen et al. 2015) especially if it is done with utmost caution and
support from conservation laws by following an extensive framework founded on ecological
and ethical considerations (Jackiw et al. 2015).

Ethics of interspecific hybridization and assisted evolution

Finally, the ethics of using interspecific hybridization is an ongoing debate. Particularly,
Filbee-Dexter & Smajdor 2019 argued that irreversible anthropogenic disturbances that have
caused the collapse of many marine and terrestrial communities are the result of our own
limited understanding of the consequences of our actions. It is therefore likely that attempting
to save reef biodiversity through assisted evolution – an approach that is both understudied and
risky – will similarly lead to further detriment of biodiversity (Filbee-Dexter & Smajdor 2019).
However, one may counterargue that it is precisely due to our current limited knowledge of
these unconventional approaches that we must keep exploring modern ways of conservation,
albeit with utmost care and consideration for the existing biodiversity. Rapidly aggravating
disturbances in the ocean warrant a continuous effort in developing a biological toolbox we
can use to aid coral reefs survive by enhancing their resilience and biodiversity (van Oppen et
al. 2015).

How interspecific hybridization can potentially increase reef resilience and overall
biodiversity.
As Chan et al. (2018) put it, “From the restoration point of view, hybridization increases genetic
variation which can potentially enhance adaptive capacity and release a population from
adaptive limits (Hoffman and Sgrò, 2011; Becker et al., 2013; Carlson et al., 2014; van Oppen
et al., 2015; Hamilton and Miller, 2016; Meier et al., 2017).” Here, inferences regarding how
coral hybrids can increase reef resilience and overall biodiversity.

It starts with choosing which coral species to hybridize. Ideally, the two species: (1) are closely
phylogenetically related to minimize pre- and post-zygotic barriers (Willis et al. 2006), (2) are
sympatric or from the same reef to prevent outbreeding (Hwang et al. 2012, Whiteley et al.
2015 as cited by Chan et al. 2018), (3) exhibit desirable traits that are heritable, so that potential
future progeny are likely to inherit them (van Oppen et al. 2015). Assuming coral hybrid
offspring is viable, successfully settles in a reef, and survives until its reproductive stage, it can
commence sexual reproduction (if it is able to produce viable gametes) either with co-hybrids
(i.e. F2 production), its parental species (i.e. backcrossing), or possibly other coral species.
This results to further gene recombination that likely contributes to increased genetic variation
and novel allele combinations that give rise to new phenotypes like adaptive capacity. Future
progenies hopefully express inherited desirable traits related to fitness and stress tolerance.
Assuming the reproduction and recombination cycle continues indefinitely, the genetic
diversity of reef-building corals likely leads to higher morphological and functional coral
diversity – creating a diverse set of microhabitats for higher organisms. The heritable fitness
and traits are likely passed onto many coral generations, thereby increasing established reef
resilience. This fitness and genetic diversity of reef-building corals ultimately translate to reef
resilience and biodiversity that sustains the overall biodiversity of the entire coral ecosystem.
This chain of events is idealized, but it demonstrates how interspecific hybridization of corals
can potentially lead to (or at least aid in achieving) enhanced reef resilience and biodiversity.

Coral hybridization experiments and resulting hybrid traits and fitness.

Biodiversity is maintained by genetic diversity of the organisms and their resilience. Resilience
is often measured through characteristics that reflect fitness and survivability despite stressors.
In coral hybridization studies, coral hybrids are considered fitter if they outperform their
purebred parents using resilience-related traits as parameters.

To make assisted evolution through hybridization work, a coral hybrid offspring must be at
least viable. The evolution of corals is reticulate, which means that divergent ancestral lineages
have converged at some point, thereby limiting pre- and post-zygotic barriers between some
coral species and allowing them to interbreed (Willis et al. 2006). This makes coral
hybridization more commonly occurring than we think, and the viability of most coral hybrids
is evidence of this. Acropora prolifera, for instance, is a naturally occurring hybrid of A.
cervicornis and A. palmata (Fogarty 2012). The hybrid of A. florida and A. intermedia on the
other hand are said to undergo normal larval development into a viable hybrid offspring
(Isomura et al. 2013).

Viable coral hybrids must have better fitness that their purebred parents to increase the
likelihood that they will be able to survive future stressors. For instance, Montastraea annularis
and M. franksi exhibit higher larvae yield compared to their purebreds. Chan et al. (2018)
showed that some Acropora hybrids (A. tenuis x A. loripes, A. sarmentosa x A. florida)
underwent normal embryonic development and had higher survival and larger recruit size than
their purebred counterparts even under high temperature and pCO2 stress. In a subsequent
study, they showed that most Acropora hybrids studied exhibited higher larval survival and
settlement even when subjected to elevated temperatures (Chan et al. 2019).

The potential of fitter coral hybrids to be acceptable in wild reef communities must also be
examined. This means not only being able to proliferate themselves, but also to not negatively
affect the native populations within the reef. For instance, a potential hybrid zone of soft coral
species Sinularia maxima and S. polydactyla was found to double its population in a decade
while populations of their purebred counterparts declined (Slattery et al. 2008). The authors
partially attributed this observation to the “deterrent” metabolites of the hybrids which were
significantly more potent that the purebreds (Slattery et al. 2008). Meanwhile, hard coral hybrid
Acropora prolifera mentioned earlier, as its name suggests, was able to proliferate through
several different reefs (Fogarty 2012, Japaud et al. 2014, Aguilar-Perera and Hernández-Landa
2017 as cited by Chan et al. 2018), even in locations wherein populations of its parental species
massively declined (Fogarty 2012). Like the soft coral, this proliferation may have been due to
its higher fitness compared to its parents (Fogarty 2012), although this greater fitness does not
necessarily deter its parental species unlike the mentioned soft coral hybrid.

Studies that examine the relative fitness of coral hybrids are still very limited, but these studies
provide evidence that coral hybridization can lead to hybrid vigor which ultimately reflects
resilience. However, higher fitness does not necessarily mean that it is not detrimental to other
wild native species (e.g., the soft coral hybrid in Slattery et al. 2008), so further examination
of coral hybrids’ potential influence on overall reef biodiversity is imperative before using said
hybrids for transplantation and reef restoration.

Recommendations on the identified key concerns/gaps

The hesitation of using interspecific hybridization in coral reef conservation largely stems from
our lack of knowledge about it, and it is therefore recommended to conduct more research on
it and how it influences the standing biodiversity and environment. Coral species that have
favorable traits and/or close phylogenetic relationships with each other are ideal subjects for
coral hybridization experiments given that there is a good chance their potential hybrid
offspring will exhibit viability and hybrid vigor. Additionally, coral hybrids deemed fitter
compared to their parental species can be examined in terms of their influences on existing reef
biodiversity.

Transplanting of said coral hybrids and their purebreds on the field can be done to monitor
relative fitness and investigate their responses to real environmental stressors (Chan et al.
2018). In relation to their effect on reef resilience and biodiversity, potential reproduction of
F1 hybrids is another aspect to explore (Chan et al. 2018). Like the production of F1 hybrids,
backcrossing can lead to another set of recombination and novel molecular interactions that
may influence the next generation’s phenotype. Exploring the viability and fitness of
subsequent generations can shed more light on the implications of having coral hybrids on the
reef (Chan et al. 2018).

The risks of interspecific hybridization remain possible despite its advantages, so it is important
to still take extra precaution in conducting coral hybridization studies. For instance, attempting
hybridization experiment is ideally done ex situ such as in a laboratory where interacting coral
gametes and ensuing hybrid larvae can be contained. Hybrid larvae for settlement must also be
separated from purebreds or wild types to avoid potentially adverse interactions among them.
Furthermore, conducting stress experiments to potential hybrids to gauge their relative fitness
is easy to do in an enclosed, controlled room. To predict their influence on other cohabiting
species, microcosm studies is also recommended before doing full-blown transplantations of
coral hybrids in wild reef communities. And if studies have reached the point where
transplantation is deemed safe, earnest planning for genetic monitoring strategies and other
considerations regarding the release of these hybrids in the wild must be done (Laikre et al.
2010). Since it is one of the parameters for choosing species to hybridize, heritability of traits
is also an aspect to be closely looked at, as well as the resultant stress response of the hybrids
to measure their relative fitness (van Oppen et al. 2015).

Finally, it must be emphasized that interspecific hybridization is not at all proposed as a

replacement for traditional methods or the sole best approach to assisted evolution.
Interspecific hybridization is but an aid to up the chances of marine life to survive the ever-
changing marine environment. It is nothing if traditional methods are not being exhausted.
Other approaches to assisted evolution are just as valid and worth of exploration as interspecific
hybridization. Thus, the best way to use interspecific hybridization as a means to enhance coral
reef resilience and biodiversity is by integrating it with other modern and already-established
conservation methods (Anthony et al. 2017).

Summary and conclusion

Coral reefs are being threatened by continuous ocean warming and acidification, and
occurrences of mass bleaching events have been more and more frequent. Due to the rapidly
changing environmental conditions, traditional conservation approaches that utilize existing
biodiversity to replenish reefs are no longer enough. As such, researchers are coming up with
new approaches to conservation such as assisted evolution which can enable enhancement of
an organism’s resilience as well as overall biodiversity. Under assisted evolution is
interspecific hybridization which occurs when two genetically distinct individuals mate and
produce a hybrid offspring that hopefully exhibits hybrid vigor. Even though hybridization
seems a promising method to enhance fitness, a lot of concerns surround it including possibility
of outbreeding depression, loss of current genetic diversity, and the ethics of its use. While all
of these are possible, they are not that likely to occur, and the need to develop a biological
toolbox to enhance reef resilience and biodiversity calls for continuous research on it. So far,
most studies on coral hybridization show increased fitness and better traits of hybrids and
further support the potential of this approach to help in increasing reef resilience and
biodiversity. Recommendations largely include further studies on coral hybridization and
taking steps towards conducting studies on their ecological implications. It is emphasized that
interspecific hybridization of corals is an aid rather than a replacement for traditional
conservation methods or other approaches to assisted evolution.

There is still much we do not know about how interspecific hybridization works or how exactly
it can enhance reef resilience or biodiversity. It remains true and is a valid concern that using
interspecific hybridization on corals can potentially be hazardous to current reef biodiversity,
but the possible benefits of it (most of it still in theory for now) is promising enough that we
must explore it – especially that current studies show that most coral hybrids are fitter than
their parental counterparts. And even if our future studies show that interspecific hybridization
is more detrimental than helpful to marine life, at least then we can say for sure and move on
to explore the next promising conservation approach.
However, we must also not rely solely on new, more intrusive methods to save marine
biodiversity. For many decades, we have employed traditional conservation efforts like habitat
restoration and MPA network establishment, and we ought to continue doing them. Again,
these methods are not proposed to completely replace traditional ones, but rather to
complement them and increase the likelihood that marine populations are able to survive.
Learning more about these approaches now will enable educated decisions regarding its use in
the future.

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