Conomurex Luhuanus
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''Conomurex luhuanus'', commonly known as the strawberry conch or tiger conch, is a
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of medium-sized
sea snail Sea snails are slow-moving marine (ocean), marine gastropod Mollusca, molluscs, usually with visible external shells, such as whelk or abalone. They share the Taxonomic classification, taxonomic class Gastropoda with slugs, which are distinguishe ...
, a marine
gastropod Gastropods (), commonly known as slugs and snails, belong to a large Taxonomy (biology), taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, freshwater, and fro ...
mollusk Mollusca is a phylum of protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000  extant species of molluscs are recognized, making it the second-largest animal phylum after Arthropoda. The ...
in the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Strombidae Strombidae, common name, commonly known as the true conchs, is a Taxonomy (biology), taxonomic family (biology), family of medium-sized to very large sea snails in the superfamily (zoology), superfamily Stromboidea, and the Epifamily Neostromboid ...
, the true
conch Conch ( , , ) is a common name of a number of different medium-to-large-sized sea snails. Conch shells typically have a high Spire (mollusc), spire and a noticeable siphonal canal (in other words, the shell comes to a noticeable point on both ...
s.Rosenberg, G. (2011). Conomurex luhuanus (Linnaeus, 1758). Accessed through: World Register of Marine Species at http://www.marinespecies.org/aphia.php?p=taxdetails&id=565370 on 2011-03-25 ''C. luhuanus'' is found in sandy habitat among corals in the Indopacific region. They feed on algae or detritus, move with a modified foot, and have complex eyes compared to other gastropods.


Shell description

The shell of ''C. luhuanus'' ranges from a more common length of 5 cm, to a maximum length of 8 cm.Poutiers, J. M. (1998). Gastropods in: FAO Species Identification Guide for Fishery Purposes: The living marine resources of the Western Central Pacific Volume 1. Seaweeds, corals, bivalves and gastropods. Rome, FAO. page 475. ''C. luhuanus'' is often mistaken for a
cone snail Cone snails, or cones, are highly venomous sea snails that constitute the family Conidae. ''Conidae'' is a taxonomic family (previously subfamily) of predatory marine gastropod molluscs in the superfamily Conoidea. The 2014 classification of ...
, mainly because of the conoidal outline of its shell, which is relatively unusual among the
Strombidae Strombidae, common name, commonly known as the true conchs, is a Taxonomy (biology), taxonomic family (biology), family of medium-sized to very large sea snails in the superfamily (zoology), superfamily Stromboidea, and the Epifamily Neostromboid ...
. Its
stromboid notch The stromboid notch is an anatomical feature which is found in the gastropod shell, shell of one Taxonomy (biology), taxonomic family of medium-sized to large sea snails, the conches. Marine (ocean), Marine gastropods in the family Strombidae hav ...
, though not as conspicuous as observed in many species that used to belong to ''
Strombus ''Strombus'' is a genus of medium to large sea snails, marine (ocean), marine gastropod molluscs in the family (biology), family Strombidae, which comprises the true conchs and their immediate relatives. The genus ''Strombus'' was named by Swedi ...
'' (such as '' Lentigo lentiginosus''), is deep and easily distinguished. The interior may be slightly lirate. The external color of the shell can be either white with an orange, brown/tan pattern of blotches, or completely brown/white. The interior is usually colored strong orange, red or pink, and the inner lip border is black or chocolate brown.


Phylogeny

The phylogenetic relationships among the Strombidae have been mainly accessed on two occasions, by Simone (2005) and Latiolais (2006), using two distinct methods. Simone proposed a
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
(a tree of descent) based on an extensive
morpho Morpho may refer to: * ''Morpho'' (genus), a genus of butterflies * Morpho (VTOL), a Swiss prototype drone * Morpho (company) IDEMIA (formerly known as OT-Morpho) is a French multinational technology company headquartered in Courbevoie, Fra ...
-
anatomical Anatomy () is the branch of morphology concerned with the study of the internal structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old scien ...
analysis of representatives of
Aporrhaidae Aporrhaidae is a family of sea snails commonly called the "pelican's foot snails." The taxonomy of the Gastropoda by Bouchet & Rocroi, 2005Bouchet P. & Rocroi J.-P. (Ed.); Frýda J., Hausdorf B., Ponder W., Valdes A. & Warén A. 2005. ''Classif ...
, Strombidae,
Xenophoridae Xenophoridae, common name, commonly called carrier shells, is a family (biology), family of medium-sized to large sea snails, marine (ocean), marine gastropod mollusks in the clade Littorinimorpha. Distribution The Xenophorids live on sand and ...
and
Struthiolariidae Struthiolariidae is a family (biology), family of small to medium-sized sea snails, marine (ocean), marine gastropod molluscs in the superfamilia, superfamily Stromboidea.Bouchet, P.; Rosenberg, G. (2014). Struthiolariidae Gabb, 1868. Accessed th ...
, which included ''C. luhuanus''. With the exception of ''
Lambis ''Lambis'' is a genus of large sea snails sometimes known as spider conchs, marine (ocean), marine gastropod mollusks in the family Strombidae, the true conch family.Bouchet, P.; Rosenberg, G. (2011)Lambis Röding, 1798 Accessed through: World Re ...
'' and '' Terebellum'', the remaining taxa were previously allocated within the genus ''
Strombus ''Strombus'' is a genus of medium to large sea snails, marine (ocean), marine gastropod molluscs in the family (biology), family Strombidae, which comprises the true conchs and their immediate relatives. The genus ''Strombus'' was named by Swedi ...
'', including ''C. luhuanus''. However, according to Simone, only '' Strombus gracilior'', ''
Strombus alatus ''Strombus alatus'', the Florida fighting conch, is a species of medium-sized, warm-water sea snail, a marine gastropod mollusk in the family Strombidae, the true conchs. Its name derives two Latin words. Strombus means, in Latin, a snail with s ...
'' and ''
Strombus pugilis ''Strombus pugilis'', common names the fighting conch and the West Indian fighting conch, is a species of medium to large sea snail, a marine gastropod mollusk in the family Strombidae, the true conchs. ''S. pugilis'' is similar in appearance t ...
'', the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
, remained within ''Strombus'', as they constituted a distinct group based on at least five
synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to ...
(traits that are shared by two or more
taxa In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
and their
most recent common ancestor A most recent common ancestor (MRCA), also known as a last common ancestor (LCA), is the most recent individual from which all organisms of a set are inferred to have descended. The most recent common ancestor of a higher taxon is generally assu ...
). The remaining taxa were previously considered as
subgenera In biology, a subgenus ( subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between the ge ...
, and were elevated to genus level by Simone in the end of his analysis. The genus '' Conomurex'', in this case, only included ''C. luhuanus''. In a different approach, Latiolais ''et al''. (2006) proposed a
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
based on sequences of nuclear
histone H3 Histone H3 is one of the five main histones involved in the structure of chromatin in eukaryotic cells. Featuring a main globular domain and a long N-terminal end, N-terminal tail, H3 is involved with the structure of the nucleosomes of the 'b ...
gene and mitochondrial cytochrome-c oxidase I (COI) gene showing phylogenetic relationships of 32 species that used to belong to the genus ''Strombus'' and ''
Lambis ''Lambis'' is a genus of large sea snails sometimes known as spider conchs, marine (ocean), marine gastropod mollusks in the family Strombidae, the true conch family.Bouchet, P.; Rosenberg, G. (2011)Lambis Röding, 1798 Accessed through: World Re ...
'', including ''Conomurex luhuanus'' (= ''Strombus luhuanus''). All species in this cladogram are still given under their original names in ''Strombus'' and ''Lambis''.


Habitat and ecology

''C. luhuanus'' dwells in sandy substrate in tropical habitats, such as the
Great Barrier Reef The Great Barrier Reef is the world's largest coral reef system, composed of over 2,900 individual reefs and 900 islands stretching for over over an area of approximately . The reef is located in the Coral Sea, off the coast of Queensland, ...
and southern Papua New Guinea. They often form colonies that move together over long periods of time. Two types of colonies have been described: colonies of juveniles that are made up of conch of the same size-class, and colonies of mixed age-classes that contain individual conch of a spectrum of ages, although within the colony separation based on age still persists. ''C. luhuanus'' are herbivores or feed on detritus. They use their
proboscis A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular arthropod mouthparts, mouthparts used for feeding and sucking. In vertebrates, a pr ...
to pick up sand which they may then swallow or eat algae from. ''Strombus'' also have sensory tentacles at the end of their eyestalks which may play a role in chemically sensing the presence of nearby food. The sensory tentacles may also be important in detecting predators, such as carnivorous snails. Unlike many snails that move by slowly creeping along their terrain, members of the Strombus family move with an awkward leaping motion as they thrust off the sea floor with their modified foot. This leaping motion is also used to quickly escape from predators.


Reproduction

''C. luhuanus'' form mating aggregations of copulating individuals were males and females compete for reproduction opportunities.


Vision

The eyes of members of the ''Strombus'' family are positioned at the ends of eye stalks and typically have a diameter of about 1.5-2.5 mm, which is large compared to their body size. The eye stalks extend from two notches in the shell. The eye of ''Strombus'' contains a spherical lens, a cornea, a pupil with a pigmented iris surrounding it, a vitreous body, and a cup-shaped retina. In contrast, some other
gastropods Gastropods (), commonly known as slugs and snails, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, freshwater, and from the land. Ther ...
only possess small open eyes containing a gelatinous substance. The retina is made up of several layers. The first layer, located adjacent to the vitreous body, is primarily composed of the processes of
photoreceptor cell A photoreceptor cell is a specialized type of neuroepithelial cell found in the retina that is capable of visual phototransduction. The great biological importance of photoreceptors is that they convert light (visible electromagnetic radiation ...
s. This layer is approximately 180 um thick. The next layer is characterized by an abundance of dark pigment granules. The following layer contains the cell bodies of different cell types, and the last layer consists of
neuropil Neuropil (or "neuropile") is any area in the nervous system composed of mostly unmyelinated axons, dendrites and glial cell processes that forms a synaptically dense region containing a relatively low number of cell bodies. The most prevalent ...
. It is from the neuropil layer that the optic nerve projects in several small branches that eventually merge to form a single nerve fiber. ''Strombus'' has almost 100,000 photoreceptors, whereas some
nudibranch Nudibranchs () are a group of soft-bodied marine gastropod molluscs, belonging to the order Nudibranchia, that shed their shells after their larval stage. They are noted for their often extraordinary colours and striking forms, and they have b ...
s merely have five. Three distinct cell types have been described in the retinas of ''C. luhuanus'', one of which is a photoreceptor, another which is a
glial cell Glia, also called glial cells (gliocytes) or neuroglia, are non-neuronal cells in the central nervous system (the brain and the spinal cord) and in the peripheral nervous system that do not produce electrical impulses. The neuroglia make up ...
, and the last which may be a second type of photoreceptor.  Only one type of visual pigment has been identified in ''C. luhuanus'', suggesting that they do not possess color vision. Electrophysiological recordings that measure electrical impulses, or action potentials, from neurons have been used to study the neural processing of visual information from the eye of ''C. luhuanus''. When eyes adapted to the dark were shown brief flashes of light, a cornea-negative potential was evoked. This response was termed the "on" response. When light was presented to the dark-adapted eye for a longer period of time, the extracellular retinal potential (ERG) recordings showed the cornea-negative potential declined until a steady-state was reached. Then, the steady-state was maintained until the light stimulus ceased and the potential returned to baseline. Finally, a third type of response that fired at the cessation of light stimulus was observed and termed the "off" response. These different waveform responses could indicate that ''C. luhuanus'' is capable of complex neural processing involving excitation, inhibition, and synaptic inhibition. The "on" response could be due to excitation of the optic nerve in response to depolarizing photoreceptors, while the decline to steady-state potential could be a sign of synaptic inhibition, and the "off" activity could indicate release from light-promoted inhibition. The light-promoted "off" potentials were found to be sensitive to anesthesia by magnesium chloride, with repetitive "off" potentials eliminated within minutes of being introduced to the magnesium chloride solution. However, the light-promoted "on" potentials were not affected by the magnesium chloride and would continue firing over an hour after being immersed in the magnesium chloride solution. These results could indicate that nerve fibers that are responsible for "off" stimuli are regulated by chemical synapses, which can be inhibited by magnesium. It is not entirely clear why ''C. luhuanus'' evolved such well-developed eyes. Usually eyes of this complexity are found on predators that move quickly through their environment looking for prey, not in slow-moving herbivorous creatures such as ''C. luhuanus''. One study suggests that the eyes are important for coordination in the escape response of ''Strombus''. When conch were blinded, they showed an increased probability of leaping nearer to the predator instead of away from it. More work remains to be done on the purpose of the eyes, however.


Regeneration

Another remarkable feature of the eyes of ''Strombus luhuanus'' is that they are capable of regeneration following amputation. The first step of regeneration following severance of the optic nerve is the formation of an eye cup as the epithelium adjacent to the cut folds inward. This occurs within a day after the optic nerve is cut. The eye cup is closed by day three, and at this stage it resembles an embryonic eye. The diameter of the eye continues to increase, and meanwhile the cells of the eye differentiate to form the different layers in the retina. When the diameter of the eye reaches about 0.8 mm (after about 15 days), the retina looks like a mature adult eye. The eye will continue to grow until it becomes fully sized, with a diameter of 2 mm, which can take several months. Size, age, sex, nutritional state, or whether the left or right eye is amputated does not have a substantial influence on the regenerative prospects of the eye. In gastropod eye regeneration, the degenerating nerve is important for the formation of the new eye. New axons leave the eye cup and bundle together to form the optic nerve. The new nerve fibers seem to be attracted to the previous optic nerve, and grow down it until they make contact with the cerebral ganglion. Electroretinograms (ERGs) have been used to monitor neuron activity in the developing eye of ''C. luhuanus'' over the course of the eye's development. The ERGs of the regenerating ''C. luhuanus'' eye grow increasingly complex as the eye matures. During the first week of regeneration, a simple "on" peak was recorded, and steady-state potentials were observed in response to prolonged illumination. When the eye reached about 0.4 mm in diameter, an "off" response was observed occasionally. At a diameter of 0.6 mm, rhythmic ERG "off" responses were detected. By 0.8 mm in diameter, the ERGs of the regenerated eye resembled that of a mature adult eye, with two distinct cornea-negative "on" potentials and rhythmic "off" potentials.


References

* Walls, J.G. (1980). ''Conchs, tibias and harps. A survey of the molluscan families Strombidae and Harpidae''. T.F.H. Publications Ltd, Hong Kong


External links

* {{Taxonbar, from=Q5162153 Strombidae Gastropods described in 1758 Taxa named by Carl Linnaeus Marine gastropods