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Summary | |
Syllis is the largest genus in Syllidae (Álvarez-Campos et al. 2015). Syllidae species usually dominate marine environments and are found worldwide (Álvaros-Campos et al. 2015). The specimen is from the genus Syllis; however, it is likely that the specimen has not been described previously. The chaetae of Syllis are key to their identification (San Martín & Wosfold 2015). The proventricle is a defining feature of Syllidae and plays a role in its reproduction, regeneration and feeding (Glasby 1993; Weidhaseet al. 2016). Syllinae use the reproductive mode of schizogamy which can be seen by the epitoke at the posterior of the specimen (Teresa, San Martín & Siddal 2012). The phylogenetic status of Syllidae and Syllis are uncertain and require further research. (Álvarez-Campos et al. 2015; Aguado & Bleidorn 2010).
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Physical Description |
Size and Colouration | |
In general, Syllidae species are small and slender (Beesley, Ross & Glasby 2000). On average, Syllidae are less than 10 mm long and 1 mm wide (Beesley, Ross & Glasby 2000). The specimen falls within this typical range (Fig. 1). Syllidae’s small size enables it to move into interstitial spaces and crevices (Martins et al. 2013). The members of the genus Syllis have cylindrical bodies (Lucas, Sikorski & San Martín 2018). Many Syllidae species are colourful due to the pigmentation of their gut, epidermis,and oocytes (Beesley, Ross & Glasby 2000). The specimen is pale orange, but its middle segments are dark yellow (Fig. 2A). Moreover, there is a dark brown marking in the middle of the specimen’s body (Fig. 2B). The dark yellow and brown are likely due to the contents of the specimen’s gut or oocytes. The specimen also has a black epitoke and blue spots on its dorsal cirri (Fig. 2A; Fig. 3). There are black lines across the specimen’s body (Fig. 2B). The epidermis of Syllidae may be smooth or papillated (Beesley, Ross & Glasby 2000). The specimen has a smooth epidermis.
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The Prostomium | |
Syllidae have a rounded prostomium which is usually wider
than long (Beesley, Ross & Glasby 2000). The prostomium of the specimen has
eyes, lateral antennae, palps, and nuchal organs (Fig. 3).
The specimen has two pairs of lensed eyes which are arranged
in a trapezium (Fig. 3). The anterior pair of eyes is larger than the posterior
pair. The lenses of the eyes would have developed from the extension of
supporting cells (Glasby 1993). Syllidae also often have a smaller anterior
pair of ocelli, however, this is not apparent in the specimen.
A pair of lateral antennae, a median antenna and palps are a
defining feature for Syllidae (Beesley, Ross & Glasby 2000). Syllinae have
separate palps and Syllis have their palps fused at the base, which can be seen
in the specimen (Fig. 3; Beesley, Ross & Glasby 2000; Lucas, Sikorski, San
Martín
2018). The specimen’s palps are longer than its prostomium (Fig. 3). The
specimen may utilise its palps as one of its methods of feeding. Palps can be
used to sort organic material from sediment (Beesley, Ross & Glasby 2000).
The nuchal organs in the specimen are not prominent (Fig. 3).
Nuchal organs are a defining feature for Syllidae and the indistinctness of the
organs is a defining trait for Syllinae (Beesley, Ross & Glasby 2000).
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Segmentation | |
The segments of the specimen have appendages. These
appendages include dorsal cirri, ventral cirri and neuropodia with protruding
chaetae (Fig. 4).
The specimen’s dorsal cirri, also referred to as tentacular
cirri, are long and appear on each segment (Fig. 5). They are one cell thick
and have blue spots on them (Fig. 3; Fig. 4). One of the defining features of Syllinae
is their long and articulated dorsal cirri (Beesley, Ross & Glasby 2000).
As is common for Syllidae, the ventral cirri are below the
neuropodia, closer to the ventral side of the specimen (Beesley, Ross &
Glasby 2000). The specimen’s ventral cirri are smaller than the dorsal cirri
and do not have spots on them (Fig. 5).
The muscles of Typosyllis antoni are arranged in a similar way in the antennae, dorsal cirri, and ventral cirri (Aguado et al.
2014). These appendages all have inner longitudinal muscle fibres that run from
the base to the end (Aguado et al. 2014) However, unlike the other appendages,
the base of the dorsal cirri has cirral muscle bundles with a basal muscular
socket (Aguado et al. 2014). Typosyllis antoni shares traits such as
chaetae shape with Syllis parturiens (Aguado et al. 2014). Due to the
shared traits of these species and Typosyllis being closely related to Syllis,
it is possible that the specimen has a similar muscular arrangement in these
appendages.
The class Polychaeta derives its name from chaetae (Merz
& Woodin 2006). The chaetae are in the ventral neuropodia. The chaetae play
a key role in the identification of Syllis species (San Martín
& Wosfold 2015). In general, Syllis have falciger compound chaetae and
thick simple chaetae (Passos Ribeiro et al. 2020). The specimen has prominent compound falciger chaetae which can be seen by the dentate blades (Fig 5). These chaetae, which have hooked tips, are
usually found in polychaetes that need to anchor themselves (Merz & Woodin
2006). This includes polychaetes that live in tubes or are symbiotic or
interstitial (Merz & Woodin 2006). Syllis are interstitial and some live-in
association with other organisms. This may be the reason why the specimen
requires these chaetae. Although the anterior and posterior have the
same type of chaetae, there are some differences between them. The anterior
chaetae have longer falcigers than the posterior chaetae (Fig. 5). Moreover, the end of
the posterior chaetae is more hooked than that of the anterior (Fig. 5).
Other key physical features in the specimen’s segmentation
include its musculature, coelomic cavities, integument, and its lack of
notopodia.
The main muscular elements of polychaetes are the circular
and longitudinal fibres in the body wall, as well as parapodial, chaetal,
oblique, diagonal and dorsoventral fibres (Tzetlin & Filippova 2005).
Polychaetes also have muscular aspects associated with their septa and
mesenteries (Tzetlin & Filippova 2005). Studies conducted on the species Syllis
gracilis found that its dorsal muscle band had at least 10 muscle bundles
next to each other and two ventral bands (Parapar et al. 2019). Moreover, it
had well developed parapodial muscular fibres, a few muscles associated with
the parapodial wall and muscles attached to the aciculae and chaetae (Parapar
et al. 2019). The muscular features of Syllis gracilis have also been
described in other Syllidae species (Parapar et al. 2019). This suggests that these
muscular elements are common in Syllidae and so the specimen is likely to have
these features.
Syllidae have fluid-filled coelomic spaces between their
body wall and gut wall (Parapar et al. 2019).
The coelomic peritoneum defines the dorsal and ventral mesenteries,
however it does not enclose the foregut which is involved in the movement of the
eversible pharynx (Parapar et al. 2019). The species Syllis gracilis has
a wide coelomic cavity which is divided by septa (Parapar et al. 2019). The
specimen is likely to share the same feature since it is in the genus Syllis.
The integument is the tough outer protective layer of an
organism. The integument of polychaetes has one epithelium layer which has
multiple cell types (Storch 1968). The epithelium layer is on top of a thin
fibrous extracellular layer (Storch 1968). The integument of the antennae and
cirri of Syllidae are distinctive to other polychaetes (Parapar et al. 2019).
The antennae and cirri have a central core of musculature and nervous tissue
which is surround by different types of integumental cells (Parapar et al.
2019). An example of one of the types of integumental cells is the gland cells
which are a long and slender corkscrew shape (Parapar et al. 2019). The
function of these integumental glands, which also occur on the dorsal and
ventral surfaces of Syllidae, is not known (Beesley, Ross & Glasby 2000).
The specimen
is lacking notopodia which is arguably an autapomorphy of Syllidae (Glasby
1993).
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Pharynx | |
In many species of the genus Syllis, the pharynx starts in
the third chaetiger which is posterior to the peristomium (Fig. 6B; San Martín
& Lopez 2003). The complex Syllidae digestive track consists of four parts:
a buccal cavity, pharyngeal tube, proventricle and ventricle (Fig. 6A; Tzetlin
& Purschke 2005).
The buccal cavity is short and leads to the longer
pharyngeal tube (Beesley, Ross & Glasby 2000).
A defining feature for the subfamily Syllinae is the
straight pharyngeal tube (Fig. 6A; Beesley, Ross & Glasby 2000). In
Syllidae, the pharyngeal tube is the non-muscular part of the pharynx and often
has a circle of papillae with sensory cells at its opening (Tzetlin &
Purschke 2005). In Syllis, this is then followed by a mid-dorsal tooth
which does not have a trepan (Lucas, Sikorski & San Martín
2018; Tzetlin & Purschke 2005). Syllidae teeth are often made of a cuticle
or sclerotised cuticle (Beesley, Ross & Glasby 2000). The pharyngeal tube
leads to the proventricle.
The proventricle is the muscular part of the pharynx and can
clearly be seen through the specimen’s body (Beesely, Ross & Glasby 2000;
Fig. 6). It has often been considered as a synapomorphy of Syllidae (Teresa,
San Martín
& Siddal 2012). However, Sphaerodoridae, Naurilinielidae and Pilargidae all
have a similar structure (Aguado, Nygren & Siddall 2007). These structures
may have formed due to convergent evolution or have originated in a distant
ancestor (Aguado, Nygren & Siddal 2007). In any event, Syllidae’s
proventricle should not be considered as a synapomorphy (Aguado, Nygren &
Siddall 2007). However, Syllidae’s proventricle is one of its defining
characteristics (Glasby 1993). The proventricle can vary greatly in length
among species (Haswell 1921). It is cylindrical and has slight lateral
compression (Haswell 1921). It is made up of striated muscles cells with sarcomeres
and calcium concretions (Weidhase et al. 2016). At the proventricle’s anterior,
there are chitinous plates which are thought to be homologous to the
sick-shaped jaws of Nereididae due to their position (Glasby 1993). The wall of
the proventricle has 6 layers: the thin splanchnic layer of the coelomic
epithelium, the thin outer fibrous membrane, the main layer composing of radial
muscle-columns and annular muscle bands, the inner fibrous membrane, the
enteric epithelium and the cuticle (Haswell 1921). As shown in figure 6A, the
proventricle has ring-shaped fine lines which are non-striated muscles fibres
and dots which are the outer ends of the cores of the radial columns of cross-straited
muscle (Haswell 1921; Tzetlin & Purschke 2005).
One role that the
proventricle plays is that it can act as a suctorial pump for feeding (Teresa,
San Martín
& Siddal 2012). It can also play a role in regeneration and the
reproductive process (Teresa, San Martín & Siddal 2012). In Syllinae, the
proventricle and prostomium work together to facilitate reproduction (Franke
1999). The proventricle releases a hormone that inhibits sexual development and
maintains the growth of segments and somatic cells (Franke 1999). This was determined by
removing the proventricle in a non-reproductive Syllinae which induced
schizogamy and then reimplanting the proventricle which stopped the schizogamy
process (Franke 1999). The prostomium hormone stimulates sexual development by
inhibiting the release of the hormone from the proventricle (Franke 1999). Moreover,
it is thought that if the hormone is above a certain level at the time of sex
determination, the gametes will be female (Heacox & Schroeder 1982). A
study on Typosyllis prolifera also found that the hormone released from
the proventricle is cyclic and high levels of it at the start of the schizogamy
period can assist with regeneration after the stolon detaches (Heacox
& Schroeder 1982). Another study that involved the removal of the proventricle
from Typosyllis antoni found that it caused schizogamy to be
accelerated, reduced the posterior segment regeneration and resulted in poor
quality stolons (Weidhase et al. 2016). This supports the idea that the
proventricle plays a key role in reproduction and subsequent regeneration. However,
since the proventricle does not have any glandular tissue, it is unclear how it
could have direct involvement in these processes (Weidhase et
al. 2016).
The ventricle of Syllidae has thin walls and is anterior to
a ciliated epithelium (Beesley, Ross & Glasby 2000). The ventricle of
Syllidae sometimes has paired water filled caeca which are thought to control
buoyancy (Tzetlin & Purschke 2005; Beesley, Ross & Glasby 2000). The
ventricle leads into the intestine and is often viewed as being a part of it (Tzetlin
& Purschke 2005).
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Epitoke | |
The specimen's epitoke formed at the posterior of its body and
has chaetae (Fig. 7.) The chaetae appear to be capillary chaetae which are for
locomotion, stabilisation and sensing the environment (Merz & Woodin 2006).
The epitoke is involved in the process of schizogamy which is a form of sexual
reproduction (Teresa, San Martín & Siddal 2012). This means that
the specimen is an adult as it can reproduce. The epitoke separates from the adult
after it has undergone morphological changes (Weidhase et al. 2017). After the
epitoke separates, the specimen will regenerate its posterior end (Weidhase et
al. 2017). In Syllinae, the epitoke may develop a head appendage before or
after it separates (Beesley, Ross & Glasby 2000). Once it has separated, it
becomes a stolon that will actively swim to the open sea to spawn (Teresa, San
Martín
& Siddal 2012). After spawning, the stolon will die (Beesley, Ross &
Glasby 2000).
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The Specimen's Identity | |
The specimen appears to be similar to the species Syllis
maganda, however there are several differences. They both have palps that are longer than the prostomium and 4 eyes in a trapezoidal arrangement (José Martínes & San Martín 2020). S. maganda is bright
orange and blue with red lines across its body and red spots on it dorsal cirri (José Martínes & San Martín
2020). The specimen is also orange, however, is not as brightly coloured. The specimen has black lines and blue spots, rather than red lines and spots. S. magnada has thin ventral cirri that are shorter than the
parapodial lobes (José
Martínes & San Martín 2020). Comparatively, the specimen’s ventral
cirri appear to be slightly longer than its parapodial lobes (Fig. 4). The
specimen has a shorter pharynx and shorter dorsal cirri but longer proventricle than S.
maganda (P Álvarez-Campos 2021, pers. comm., 22 May). Moreover, the
specimen’s anterior and posterior chaetae have shorter blades and its posterior chaetae also have thinner shafts than those of S. maganda (Fig. 6; Fig.
8; P Álvarez-Campos
2021, pers. comm., 22 May). Given the differences to S. maganda,
the specimen is likely to be a new species (P Álvarez-Campos 2021, pers. comm., 22
May).
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Figure 1 |
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Figure 2 |
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Figure 3 |
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Figure 4 |
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Figure 5 |
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Figure 6 |
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Figure 7 |
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Figure 8 |
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Ecology |
Habitat | |
Syllidae are common in Australian waters and usually
dominant marine habitats in terms of number of species and individuals (Álvarez-Campos
et al. 2015). The success of Syllidae is likely due to their wide range of body
sizes, feeding mechanisms and reproductive strategies (Serran, San Martín &
López 2006). The genus Syllis is the largest and most diverse genus in Syllidae
(Álvarez-Campos et al. 2015). Syllis has more than 120 species globally and 30
species in Australia (Álvarez-Campos et al. 2015). The abundance of Syllis species depends on the type of environment. For instance, Syllis
armillaris and Syllis variegata dominate subtidal areas, whereas, Syllis gracilis
dominates
intertidal and shallow subtidal areas as it is suited to less structured
environments (Serrano, San Martín & López 2006).
Syllidae are found in areas with soft substrates where they generally have an
interstitial life (Granados-Barba et al. 2003; San Martín & Worsfold 2015).
They are also found in areas with hard substrates where they have an errant life and live
among organisms such as algae, as well as biogenic structures (Granados-Barba
et al. 2003; San Martín & Worsfold 2015). Syllidae are especially abundant
in shallow marine habitats but are also found in deeper waters (San Martín
& Worsfold 2015). They are not found in fresh water and do not play an
important role in estuaries (San Martín & Worsfold 2015). Syllidae are also
found in cryptic environments, such as sponges and dead corals, where they generally
have the highest species richness (Granados-Barba et al. 2003). For example, Syllis
mayeri is associated with the sponge Ircinia strobilina (Musco &
Giangrande 2005). Additionally, Syllidae are found in biofouling communities on
artificial substrates (Diaz-Castaneda 2000). The presence of seaweed and other
invertebrates may assist with the colonisation of Syllidae because they prefer
holes and crevices (Ba-Akdah 2018). However, studies have found that Syllidae are
one of the dominant groups in the early stages of colonisation which suggests
that they do not need the assistance of other organisms to colonise an area (Diaz-Castaneda 2000).
Another study found that Syllis species were some of the first to colonise an
artificial area (Ba-Akdah 2018). The ability of Syllis species to colonise
artificial substrates explains why the specimen was present on the ARMS
plate that was hung off One Mile Jetty in Dunwich, North Stradbroke Island from
the 27th of January to the 24th of March.
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Feeding | |
Most species in Syllinae are carnivorous (Franke 1999).
There is a lack of consensus on whether most Syllidae are generalist or
specialist feeders (Martins et al. 2013; Franke 1999). The argument for Syllidae
being generalist feeders is that they are active and live amongst a wide range
of organisms (Martins et al. 2013). This means that Syllidae have many
different preys available to them and they can move around to obtain their
food. An example of a generalist Syllis species is Syllis gracilis (Serrano,
San Martín
& López
2006). It is likely that Syllidae feed on corals, hydrozoans, bryozoans, sponges,
and ascidians (Franke 1999; Beesley, Ross & Glasby 2000).
Syllis feed through the use a few structures. First, the
pharyngeal tooth is used to pierce prey (Beesley, Ross & Glasby 2000). It
has been suggested that the pharyngeal tooth also has a basal poison gland
(Beesley, Ross & Glasby 2000). Secondly, the proventricle acts as a pump to
suck up the contents of the prey (Haswell 1921; Beesley, Ross & Glasby 2000)
Thirdly, the chitinous plates in the proventricle may facilitate feeding by
mechanically digesting food (Haswell 1921).
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Contributions to the Environment | |
Given the abundance of Syllis species, they play an important role in
the environment. Syllidae live in association with many organisms and have relationships that range from symbiotic to parasitic (López et al. 2001). Symbiotic
relationships benefit both parties, whereas parasitic relationships negatively
impact the host. Despite the negative implications for the host, parasitic relationships are still important as
they can affect the food web and interactions in the environment. As
established prior, Syllidae are good at colonising empty spaces, which means
that they would facilitate the colonisation of other species (Diaz-Castaneda
2000). Syllidae are predated on by marine benthic organisms and provide food
for pelagic organisms when their stolons migrate to the pelagic environment (Ertan
Cinar 2003). Syllinae are also predators and so play a key role in population management
(Ertan Cinar 2003).
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Life History and Behaviour |
Life History | |
Syllidae can reproduce sexually and asexually (Beesley, Ross
& Glasby 2000). In general, species have separate sexes, however there are
some Syllinae species that are hermaphrodites (Beesley, Ross & Glasby 2000).
Asexual reproduction is not very common in Syllidae because usually
the posterior cannot replace the pharyngeal system (Franke 1999). Syllis
gracilis is one of the few species that can completely regenerate and so is
able to reproduce asexually (Franke 1999). Since Syllis gracilis is from
the same genus as the specimen, it is possible that the specimen can asexually
reproduce.
Sexual reproduction in Syllidae occurs by epitoky which
involves the adult undergoing metamorphosis before spawning (Ribeiro, Bleidorn
& Aguado 2018). There are two types of epitoky: epigamy and schizogamy,
however Syllinae only undergo schizogamy (Fig 9; Ribeiro, Bleidorn & Aguado
2018; Teresa, San Martín & Siddal 2012). This is a very successful mode of
reproduction in Syllinae (Franke 1999).
The process of schizogamy begins when the atokous form of
Syllinae metamorphoses into an epitokous form (Fig 9(2)). The specimen is in
this stage of its lifecycle as it has an epitoke (Fig. 7). The epitoke,
which is in the posterior segments of the body, is detached as a stolon
(Teresa, San Martín & Siddal 2012). The adult must then regenerate its
posterior segments (Fig 9(7)). The stolons in Syllinae do not display sexual
dimorphism, however, there is variation in stolon structure (Franke 1999).
There are five types of stolons in Syllinae. First, acephalous where the stolon
has no anterior appendages (Teresa, San Martín & Siddal 2012). Secondly,
acerous where the stolon has two pairs of eyes (Teresa, San Martín
& Siddal 2012). Thirdly, dicerous where the stolon has two pairs of eyes and
a pair of unarticulated antennae (Teresa, San Martín & Siddal 2012). Fourthly,
tetracerous where the stolon has two pairs of eyes and a pair of well-developed
antennae (Teresa, San Martín & Siddal 2012). Fifthly, pentacerous where the stolon
has two pairs of eyes, a pair of lateral articulated antennae and a central
antenna (Teresa, San Martín & Siddal 2012). The specimen has an acephalous
stolon as it does not have any eyes or appendages (Fig. 7). Moreover, Syllinae
have sexual lability where individuals can produce stolons of different sexes
in a subsequent schizogamy event (Franke 1999).
Once the stolon is detached, it actively swims to the
pelagic environment (Fig 9(3); Teresa, San Martín & Siddal 2012). Broadcast
spawning occurs when Syllinae stolons discharge eggs of about 70 μm and sperm
(Fig 9(4); Franke 1999). After releasing eggs or sperm, the stolon dies (Fig
9(5)). The eggs then sink to the bottom and within 24-48 hours, the larvae
hatch as trochophore (Franke 1999). Despite having small eggs, Syllinae larvae
have a small pelagic phase which only lasts a few hours or days (Franke 1999).
This is likely due to the fact that the epitoke already provides some dispersal
in the pelagic environment and it is important that the larvae do not disperse
too far as Syllinae only have a few suitable habitats (Franke 1999).The larvae
then move into the metatrochophore stage where the larvae swim near the bottom
and become completely benthic (Franke 1999). The larvae develop into juveniles
and then adults (Fig 9(6)).
Another potential reproductive strategy that the specimen
may employ is the one of Syllis rosea which involves the female stolon taking
care of the offspring (Langeneck et al. 2020). The female stolon releases the
eggs in a gelatinous cluster which eventually compacts into a cocoon and is attached
to the middle-posterior chaetigers (Langeneck et al. 2020). The larvae remain
associated with the female stolon until it dies 7 days later (Langeneck et al.
2020). Although this is not a common reproductive strategy, it occurs within
the specimen's genus and so the specimen potentially uses this method.
The initiation of schizogamy has many potential causes. The
external and internal factors that might cause the commencement of schizogamy
are thought to target the proventricular and prostomial hormones (Franke 1999).
The first potential trigger is the season. During summer, the prostomium goes
through short recurring phases of high hormone release and as a response, the
proventricular endocrine activity reduces (Franke 1999). This results in reproductive
phases throughout summer. The specimen was collected from Dunwich on the 24th
of March. The temperature around the time the specimen was collected was above the temperature used in an experiment that found summer temperatures could trigger reproduction (Franke 1999). The specimen's reproduction could have been caused by it being summer. The second potential trigger is light. It
has been found that in Typosyllis prolifera the photoperiod
determines whether reproduction occurs and a temperature of around 13˚C
is a mere prerequisite (Franke 1999). Experiments in Syllis amica also
found that photoperiod was important in triggering reproduction and so this may
be a trigger for the specimen too (Franke 1999). The third potential trigger is
the phases of the moon, however there is no conclusive evidence in the genus
Syllis to support this idea (Franke 1999). The fourth potential trigger is
night-time as it has been found that swarming tends to occur in the evening for
Syllis amica due to the tidal cycle and so this is also a
potential trigger for the specimen (Franke 1999). This trigger is likely to be a strategy
to reduce predation risk (Franke 1999). The fifth potential trigger is a
pheromonal interaction between swarming males and females (Franke 1999).
However, there has been no evidence of any Syllis species undergoing
reproduction due to a pheromonal interaction and so this is unlikely to be a
trigger for the specimen.
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Figure 9 |
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Behaviour | |
33 species of Syllidae live in close association with other marine
invertebrates (López et al. 2001). The commensal Syllidae species are associated
with sponges, cnidarians, sipunculids, echinoderms, bryozoans, and decapods (López
et al. 2001). The parasitic Syllidae species are associated with sponges but
can also be parasites of cnidarians, other polychaetes, nemerteans, and
tunicates (López
et al. 2001).
An example of Syllis species that live in association other
invertebrates are Syllis cf. armillaris, Syllis ferrani, and Syllis
pontxioi (López et al. 2001). They live inside gastropod shells which
are occupied by hermit crabs and Pionosyllis magnicca (López
et al. 2001). These Syllis species do not have any morphological adaptations to
this mode of life and have been reported as free-living as well as living in
association with colonial or reef-building invertebrates (López
et al. 2001). More research is required to determine the nature of the association
and whether these species have any behavioural adaptations (López
et al. 2001).
Syllis species have also been found to host protozoans (Álvarez-Campos
et al. 2014). Protozoans were found on the parapodial bases of Syllis magdalena
and Syllis prolifera (Álvarez-Campos et al. 2014). They were also
found on the dorsal surface, nuchal organs, mouth opening and anterior cirri of
Syllis elongata (Álvarez-Campos et al. 2014)
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Anatomy and Physiology |
Regeneration | |
Generally, Syllidae regenerate from the pygidium, prostomium or midbody
segments (Ribero, Bleodorn & Aguado 2018). However, many Syllinae species
demonstrate the ability to regenerate their anterior (Ribero, Bleodorn &
Aguado 2018). This increased regenerative ability is linked to their use of schizogamy
which has a regenerative aspect (Weidhase et al. 2017). The most common method
of regeneration in Syllidae is an epimorphic process, however some species use
morphallaxis (Ribero, Bleodorn & Aguado 2018). Epirmorphosis involves somatic
cell activity, dedifferentiation, and re-differentiation (Ribero, Bleodorn
& Aguado 2018). Dedifferentiation involves the formation of a blastema that
contains undifferentiated cells and acts as a growth zone (Ribero, Bleodorn
& Aguado 2018). Comparatively, morphallaxis does not involve blastema
formation and so the body remodels itself so that the morphology matches the
role of the new segment position (Ribero, Bleodorn & Aguado 2018). Regeneration
time varies between species (Ribero, Bleodorn & Aguado 2018).
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Excretory System | |
Syllis has a metanephridial system for excretion of waste products. As in Syllis gracilis, the specimen likely has metanephridia with metameric disposition (Parapar et al. 2019). Metanephridia occur in all body segments except for the most anterior and posterior ones (Goodrich 1945). The metanephridial duct opens externally via a ventral nephridiopore and internally into the coelom of the anterior segment by a nephrostome (Beesley, Ross & Glasby 2000). According to Goodrich (1945), in species with stolons, such as Syllis species, the metanephrida and gonoducts fuse to form a metanephromixia. However, Bartolomaeus (1999) does not think that that the metanephromixia is formed by the fusion of two types of tissues but rather from just one tissue. In any event, the metanephromixia is involved in excretion and the release of gametes (Parapar et al. 2019).
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Nervous System | |
The general features of an Annelida central nervous system can be observed in Syllis gracilis and so are likely to be present in the specimen (Parapar et al. 2019). The Annelida central nervous system consists of an anterior prostomial brain, ventral nerve cord, segmentally arranged ganglia and circumpharyngeal nerves (Orrhage & Müller 2005). Capillary chaetae are also thought to play a role as mechano-receptors (Merz & Woodin 2006).
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Circulatory System | |
Polychaetes have a closed circulatory system which usually
consists of capillary beds, gut lacuna, and medial, dorsal, and ventral longitudinal
vessels that are linked by smaller vessels (Beesley, Ross & Glasby 2000).
The blood flows towards the posterior in the ventral vessel which is below the
gut and has branches that link to the body wall muscles and epidermis (Beesley,
Ross & Glasby 2000). The blood then flows through the lateral vessel to the
anteriorly flowing dorsal vessel, after which the blood goes around the gut and to
the ventral vessel again (Beesley, Ross & Glasby 2000). Body wall movements
and muscles that surround the dorsal blood vessel control blood flow (Beesley,
Ross & Glasby 2000).
The circulatory system of Syllis gracilis is less
complex than other polychaetes (Parapar et al. 2019). S. gracilis has two
major dorsal and ventral blood vessels surrounded by coelomic peritoneum, as
well as thin vessels that connect the two major longitudinal vessels (Parapar
et al. 2019). Moreover, the circulatory system of S. gracilis appears to
be less complex than Syllis garciai as it lacks a well-developed pair of
longitudinal lateral vessels on either side of the gut (Parapar et al. 2019).
This suggests that the circulatory systems in Syllis are likely to vary between
species. Therefore, the exact nature of the circulatory system in the specimen
cannot be determined without further examination. However, it is likely that the specimen has a simple circulatory system.
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Respiratory System | |
Due to the size of Syllidae, they generally do not possess
respiratory organs (Purschke et al. 2017). Instead, it is likely that the specimen
has respiratory pigments dissolved in its blood that transport oxygen to its
tissues (Beesley, Ross & Glasby 2000). In general, polychaetes can draw
50-60% of oxygen from the water that touches their body (Beesley, Ross &
Glasby 2000).
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Digestive System | |
The digestive system of polychaetes includes a foregut, midgut, and hindgut (Tzetlin & Purschke 2005). Despite the metameric segmentation in Syllidae, its foregut is highly specialised (Haswell 1921). The foregut includes a pharyngeal sheath, pharyngeal tube, proventricle, ventricle, and lateral caeca (Haswell 1921). These features are described in the pharynx section. The midgut of polychaetes is adapted to the species’ needs and consists of a stomach and intestine (Tzetlin & Purschke 2005). It is likely that the intestine would not be longer than the specimen’s body and would not be coiled (Tzetlin & Purschke 2005). Septa and mesenteries attach the gut system to the body wall (Tzetlin & Purschke 2005). Further examination of the specimen would be required to find out about its midgut and hindgut.
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Biogeographic Distribution | |
Some species of Syllis are found worldwide (Álvaros-Campos et al.
2015). Species have been described in Australia, Brazil, Canary Islands,
Hawaii, Indonesia, Japan, Mediterranean Sea, New Zealand, Philippines, and
Polynesia (Matos Nogueira & San Martín 2002; Álvaros-Campos et al. 2015). Matos
Nogueira and San Martín (2002) expressed some apprehension in extending the
location of Syllis from the Pacific Ocean to the Atlantic, however, they could
not find any significant distinguishing features between their specimens and
those in the Pacific Ocean. In Australia, Syllis have been identified in New South Wales, the Northern Territory, Queensland, Victoria, and Western Australia (Álvaros-Campos
et al. 2015). Moreover, Syllis are in Dunwich, North Stradbroke Island
as this is where the specimen was found.
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Evolution and Systematics |
Phylogenetic Relationships of Syllidae and Syllinae | |
The monophyly of Syllidae has been supported by the incorporation
of new taxa and the secondary structure of 18S rDNA (Teresa, San Martín
& Siddal 2012). Moreover, the subfamilies of Syllidae are all monophyletic,
except for Eusyllinae (Aguado, Nygren & Siddall 2007).
The families Calamyzidae and Levidoridae are no longer valid
and are now considered to be members of Syllidae (Beesley, Ross & Glasby
2000). Comparatively, Phyllodocida used to be considered a clade, however, is
now classified as an order (Fauchald 1977).
Syllidae was suggested to be part of the superfamily Nereidoidea,
however this was challenged by the fact that Nereidoidea may be paraphyletic
(Beesley, Ross & Glasby 2000). It has been proposed that Syllidae is part of new superfamily (Aguado & Bleidorn 2010).
This proposition stems from the idea that Syllidae evolved rapidly which led to
the creation of two ancient evolutionary lines (Aguado & Bleidorn 2010). The
first consists of Anoplosyllinae and is characterised by its chaetal and dorsum
features (Aguado & Bleidorn 2010). The second includes the rest of the Syllidae
and is characterised by an increased rate of evolution in the 18S molecule (Aguado
& Bleidorn 2010). The superfamily, Sylloidea, and two families, Anoplosyllidae
and Syllidae, have been proposed (Aguado & Bleidorn 2010).
Syllidae reproduction provides important phylogenetic information
(Garwood 1991; Franke 1999). Epigamy is thought to be the primitive condition
and schizogamy the derived condition (Teresa, San Martín & Siddall 2012). Schizogamy
is hypothesised to have evolved independently in Syllinae and Autolytinae
(Franke 1999).
A study was conducted to compare the muscular systems of
Syllidae and Sphaerodoridae (Filippova et al. 2010). It was found that despite the
families’ proventricles being similar, there were differences between the
structures, including their fibre composition (Filippova et al. 2010). It was concluded
that the families were not a sister group and that the proventricle is a homoplasy
of the families (Filippova et al. 2010; Teresa, San Martín & Siddal 2012).
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Phylogenetic Relationships of Syllis | |
There is a lot of uncertainty surrounding the phylogenetic
relationships of Syllis (Álvarez-Campos et al. 2015). The first reason for this is because
Syllis has no clear synapomorphies and was traditionally thought of as a genus
with a mixture of species that had no distinctive features (Álvarez-Campos
et al. 2015). The second reason is that there is a lack of detail in the old descriptions
of Syllis species, the old specimens have been lost and there is a lack of
molecular data for many species (Álvarez-Campos et al. 2015).
In 1879, Langerhans divided Syllis into four subgenera: Typosyllis
which have falcigerous chaetae, Haplosyllis which have thick simple chaetae, Ehlersia
which have falcigerous and elongated compound chaetae, and Syllis which have pseudo-simple
and falcigerous chaetae (Álvarez-Campos et al. 2015). Haplosyllis is now considered
a distinct genus in Syllinae, however, the placement of the other groups of species
is debatable (Álvarez-Campos et al. 2015).
Licher (1999) considered Typosyllis and Syllis to be
separate genera (Álvarez-Campos et al. 2015). It was suggested that Typosyllis had
213 taxa, including the Ehlersia species (Álvarez-Campos et al. 2015). Licher (1999)
thought that only species with pseudo-simple chaetae could be Syllis (Álvarez-Campos
et al. 2015). However, Aguado et al. (2014) found that this analysis was
inconsistent with the relationships between the species. This is because simple
chaetae evolved several times by the fusion of shafts and blades, and other
species also evolved simple chaetae by the loss of blades (Aguado et al. 2014).
Moreover, San Martín (2003) thought that Syllis was the only valid genus (Álvarez-Campos
et al. 2015). Some of the Syllis and
Typosyllis species in Australia were synonymised, sometimes due to error, which
has created more confusion surrounding the classification of these species (Álvarez-Campos
et al. 2015). One phylogenetic study suggested that Syllis and Typosyllis
are paraphyletic, however, the results were inconclusive (Álvarez-Campos et
al. 2015). If future research supported these results, it is likely that Typosyllis
would become invalid (Álvarez-Campos et al.
2015).
To resolve the uncertainty surrounding Syllis and Typosyllis, more
information about stolons, molecular data and sampling is
required (Teresa, San Martín & Siddal 2012; Aguado et al. 2014). Until
further clarification, the original classification of species as Syllis or
Typosyllis may be used (Passos Ribeiro et al. 2010).
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Taxonomic Classification | |
Kingdom: Animalia
Phylum: Annelida
Class: Polychaeta
Order: Phyllodocida
Family: Syllidae
Subfamily: Syllinae
Genus: Syllis
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Conservation and Threats | |
Since Syllidae are usually the dominant group in marine habitats, they are not endangered (Álvarez-Campos et al. 2015). Moreover, Syllis is not endangered either as it is the largest group in Syllidae (Álvarez-Campos etal. 2015). The specimen is probably not under threat given the success of other members in its genus.
Although Syllis is not endangered, there are many threats
that could cause harm to Syllis. One potential threat is habitat loss (Gray
1997). For instance, it was estimated that most of Southeast Asia’s reefs will
be destroyed within 40 years (Gray 1997). Although Syllidae are found in dead
corals, the destruction of reefs would harm Syllidae. This is because the
species that Syllidae relies on in coral habitats would die, the dead coral
would eventually disappear, and this habitat would no longer exist. The
destruction of Syllidae’s habitats would result in the loss of Syllidae species
as many would be unable to find or adapt to a new habitat. Another issue is the
fragmentation of habitats which results in increased extinctions; however, this
issue requires more research (Gray 1997). Additionally, climate change is
likely to alter storm events and rainfall which would affect the transportation
of nutrients to coasts and would have huge impacts on Syllidae in these environments
(Gray 1997). Ocean temperatures rising due to climate change not only
affects Syllidae species but also the species that they rely on (Gray
1997). Moreover, human interference with the Syllidae environments could have
serious consequences. For instance, one of the places in which trawling occurs
is in the Atlantic Ocean where some Syllidae species live (Gray 1997). Trawling
destroys habitats and organisms that live there (Gray 1997). Another example is
human trampling which can have severe consequences on coastal communities
(Plicanti et al. 2016). Thus, although Syllis is not endangered, there are many
threats which are currently harming Syllis species and could further harm them
in the future.
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References |
Acknowledgments | |
Thank you, Patricia Álvarez-Campos, Greg Rouse, and Guillermo San Martín for assisting me with the identification of the specimen and
providing me with lots of useful information. Thank you, Pat Hutchings, for
suggesting the use of POLiKEY for the identification of the family.
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