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''Titanostrombus galeatus'', the Eastern Pacific giant 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 large
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 conchs and their allies.MolluscaBase eds. (2020). MolluscaBase. Titanostrombus galeatus (Swainson, 1823). Accessed through: World Register of Marine Species at: http://www.marinespecies.org/aphia.php?p=taxdetails&id=1429806 on 2020-04-02 It is an eastern
Pacific The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean, or, depending on the definition, to Antarctica in the south, and is bounded by the cont ...
species that occurs from the
Gulf of California The Gulf of California (), also known as the Sea of Cortés (''Mar de Cortés'') or Sea of Cortez, or less commonly as the Vermilion Sea (''Mar Vermejo''), is a marginal sea of the Pacific Ocean that separates the Baja California peninsula from ...
to
Peru Peru, officially the Republic of Peru, is a country in western South America. It is bordered in the north by Ecuador and Colombia, in the east by Brazil, in the southeast by Bolivia, in the south by Chile, and in the south and west by the Pac ...
. The shells of this species were used as a
wind instruments A wind instrument is a musical instrument that contains some type of resonator (usually a tube) in which a column of air is set into vibration by the player blowing into (or over) a mouthpiece set at or near the end of the resonator. The pitch ...
by the Chavín, an ancient
civilization A civilization (also spelled civilisation in British English) is any complex society characterized by the development of state (polity), the state, social stratification, urban area, urbanization, and symbolic systems of communication beyon ...
from the northern
Andean The Andes ( ), Andes Mountains or Andean Mountain Range (; ) are the longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range is long and wide (widest between 18°S ...
highlands of Peru. They are still used for this purpose in the Andes, where they are known as ''pututu''.


Distribution

''Titanostrombus galeatus'' occurs in several countries and regions along the coastal waters of the eastern
Pacific Ocean The Pacific Ocean is the largest and deepest of Earth's five Borders of the oceans, oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean, or, depending on the definition, to Antarctica in the south, and is ...
, including the Gulf of California,
Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...
, Pacific
Panama Panama, officially the Republic of Panama, is a country in Latin America at the southern end of Central America, bordering South America. It is bordered by Costa Rica to the west, Colombia to the southeast, the Caribbean Sea to the north, and ...
,
Nicaragua Nicaragua, officially the Republic of Nicaragua, is the geographically largest Sovereign state, country in Central America, comprising . With a population of 7,142,529 as of 2024, it is the third-most populous country in Central America aft ...
,
Costa Rica Costa Rica, officially the Republic of Costa Rica, is a country in Central America. It borders Nicaragua to the north, the Caribbean Sea to the northeast, Panama to the southeast, and the Pacific Ocean to the southwest, as well as Maritime bo ...
,
Ecuador Ecuador, officially the Republic of Ecuador, is a country in northwestern South America, bordered by Colombia on the north, Peru on the east and south, and the Pacific Ocean on the west. It also includes the Galápagos Province which contain ...
, Galapagos Islands and Peru.


Description

The shell of ''Titanostrombus galeatus'' is large (maximum size 23 cm (8.97 inches), common to 14 cm (5.46 inches)), very thick and heavy. It has an oblong outline, with a short pointed
spire A spire is a tall, slender, pointed structure on top of a roof of a building or tower, especially at the summit of church steeples. A spire may have a square, circular, or polygonal plan, with a roughly conical or pyramidal shape. Spire ...
that lacks spines and nodules and is often eroded. The
body whorl The body whorl is part of the morphology (biology), morphology of the gastropod shell, shell in those gastropod mollusks that possess a coiled shell. The term is also sometimes used in a similar way to describe the shell of a cephalopod mollusk ...
is very inflated, with numerous spiral ridges and low, slightly noticeable nodules on the shoulder. The
periostracum The periostracum ( ) is a thin, organic coating (or "skin") that is the outermost layer of the shell of many shelled animals, including molluscs and brachiopods. Among molluscs, it is primarily seen in snails and clams, i.e. in gastropods an ...
is thick. The outer lip is very flared, posteriorly expanded, not higher than the apex of the spire. The edge of the outer lip edge bears a shallow
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 ...
that is often associated with the undulations originating from the superficial spiral sculpture. The
columella Lucius Junius Moderatus Columella (, Arabic: ) was a prominent Roman writer on agriculture in the Roman Empire. His in twelve volumes has been completely preserved and forms an important source on Roman agriculture and ancient Roman cuisin ...
is smooth with a well-developed callus in this species. The shell is colored ivory white to light brown externally, with a darker spire and a brown periostracum. The
aperture In optics, the aperture of an optical system (including a system consisting of a single lens) is the hole or opening that primarily limits light propagated through the system. More specifically, the entrance pupil as the front side image o ...
is bright white, and the outer lip and columellar callus are often extensively orange or dull brown in old specimens.


Phylogeny

In 2006, Latiolais and colleagues 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 ...
that attempts to show the
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
relationships of 34 species within the family Strombidae. The authors analysed 31 species in the genus ''Strombus'' including ''Lobatus galeatus'' (there referred to as ''Strombus galeatus''), and three species in the allied genus ''
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 ...
''. The cladogram was based on
DNA Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...
sequences of both 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 ...
and mitochondrial cytochrome-c oxidase I protein-coding
gene In biology, the word gene has two meanings. The Mendelian gene is a basic unit of heredity. The molecular gene is a sequence of nucleotides in DNA that is transcribed to produce a functional RNA. There are two types of molecular genes: protei ...
regions. In this proposed phylogeny, ''Strombus galeatus'' is show as a
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
to the
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
that includes ''
Lobatus gallus ''Aliger gallus'', previously known as ''Strombus gallus'', common name the rooster conch or rooster-tail conch, is a species of medium-sized sea snail, a marine (ocean), marine gastropod mollusk in the family (biology), family Strombidae, the t ...
'' (''Strombus gallus''), ''
Lobatus gigas ''Aliger gigas'', originally known as ''Strombus gigas'' or more recently as ''Lobatus gigas'', commonly known as the queen conch, is a species of large sea snail, a marine gastropod mollusc in the family of true conches, the Strombidae. This s ...
'' (''Strombus gigas''), ''
Lobatus costatus ''Macrostrombus costatus'', formerly known as ''Strombus costatus'' and ''Lobatus costatus'', or commonly known as the milk conch, is a species of large sea snail, a marine gastropod mollusk in the family Strombidae, the true conchs. They are an ...
'' (''Strombus costatus''), ''
Lobatus raninus ''Lobatus raninus'', common name the hawk-wing conch, is a species of medium to large sea snail, a marine (ocean), marine gastropod mollusk in the family (biology), family Strombidae, the true conchs.Bouchet, P. (2010). Lobatus raninus (Gmelin, 1 ...
'' (''Strombus raninus''), '' Lobatus peruvianus'' (''Strombus peruvianus'') and their
last 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 ...
.


Ecology

''Titanostrombus galeatus'' dwells on rocky, sandy bottoms near
mangrove A mangrove is a shrub or tree that grows mainly in coastal saline water, saline or brackish water. Mangroves grow in an equatorial climate, typically along coastlines and tidal rivers. They have particular adaptations to take in extra oxygen a ...
areas. It lives in depths from the
low tide Tides are the rise and fall of sea levels caused by the combined effects of the gravitational forces exerted by the Moon (and to a much lesser extent, the Sun) and are also caused by the Earth and Moon orbiting one another. Tide tables c ...
mark to 15 m, but records go as deep as 30 m. During the beginning of the year, in the early months, ''L. galeatus'' exhibits a
gregarious Sociality is the degree to which individuals in an animal population tend to associate in social groups (gregariousness) and form cooperative societies. Sociality is a survival response to evolutionary pressures. For example, when a mother was ...
behavior, forming large agglomerates in shallow waters. This is due to the
mating In biology, mating is the pairing of either opposite-sex or hermaphroditic organisms for the purposes of sexual reproduction. ''Fertilization'' is the fusion of two gametes. '' Copulation'' is the union of the sex organs of two sexually repr ...
season, which is also when
oviposition The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typica ...
occurs.
Females An organism's sex is female (symbol: ♀) if it produces the ovum (egg cell), the type of gamete (sex cell) that fuses with the male gamete (sperm cell) during sexual reproduction. A female has larger gametes than a male. Females and male ...
usually lay eggs arranged in clusters called egg masses, either directly on the sand or on dead
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 ...
shells. During the 19th century, strombid gastropods were believed to be
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they ar ...
s. This erroneous conception was based on the writings of the French naturalist
Jean Baptiste Lamarck Jean-Baptiste Pierre Antoine de Monet, chevalier de Lamarck (1 August 1744 – 18 December 1829), often known simply as Lamarck (; ), was a French naturalist, biologist, academic, and soldier. He was an early proponent of the idea that biologi ...
, whose classification scheme grouped strombids with the carnivorous sea snails. However, subsequent studies have refuted the concept completely, proving without a doubt that strombid gastropods are
herbivorous A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat n ...
animals. ''L. galeatus'' feeds primarily on
macroalgae Seaweed, or macroalgae, refers to thousands of species of Macroscopic scale, macroscopic, Multicellular organism, multicellular, ocean, marine algae. The term includes some types of ''Rhodophyta'' (red), ''Brown algae, Phaeophyta'' (brown) and ...
, and occasionally on
detritus In biology, detritus ( or ) is organic matter made up of the decomposition, decomposing remains of organisms and plants, and also of feces. Detritus usually hosts communities of microorganisms that colonize and decomposition, decompose (Reminera ...
. The giant conch is preyed upon by
invertebrates Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordate subphylum ...
, such as
octopuses An octopus (: octopuses or octopodes) is a soft-bodied, eight-limbed Mollusca, mollusc of the order (biology), order Octopoda (, ). The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttlefish, ...
, and also by
vertebrates Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
, including
rays Ray or RAY may refer to: Fish * Ray (fish), any cartilaginous fish of the superorder Batoidea * Ray (fish fin anatomy), the bony or horny spine on ray-finned fish Science and mathematics * Half-line (geometry) or ray, half of a line split at an ...
(genera and species of
cartilaginous fish Chondrichthyes (; ) is a class of jawed fish that contains the cartilaginous fish or chondrichthyans, which all have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fish'', which have skeleto ...
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 ...
Myliobatidae), triggerfish (family Balistidae), and snappers (
perciform Perciformes (), also called the Acanthopteri, is an order or superorder of ray-finned fish in the clade Percomorpha. ''Perciformes'' means "perch-like". Among the well-known members of this group are perches and darters (Percidae), and also sea ...
fish in the family
Lutjanidae Lutjanidae or snappers are a family of perciform fishes, mainly marine but with some members inhabiting estuaries and, in some cases, fresh water (e.g., '' Lutjanus goldiei''). The family includes about 113 species. Most species are used for foo ...
).


Behavior

Compared to other gastropods, ''Titanostrombus galeatus'' has an unusual means of locomotion, which is common only among the Strombidae. This curious series of maneuvers was originally described by the American zoologist
George Howard Parker George Howard Parker (December 23, 1864 – March 26, 1955) was an American zoologist. He was a professor at Harvard, and investigated the anatomy and physiology of sense organs and animal reactions. Biography George Howard Parker was born in Phi ...
in 1922. The animal initially fixes the posterior end of the foot by thrusting the point of its sickle-shaped operculum into the
substrate Substrate may refer to: Physical layers *Substrate (biology), the natural environment in which an organism lives, or the surface or medium on which an organism grows or is attached ** Substrate (aquatic environment), the earthy material that exi ...
. Then it extends its foot forward, lifting the shell and throwing it ahead in a motion that Parker called "leaping". ''L. galeatus'' is known to move long distances, on a scale of
kilometers The kilometre ( SI symbol: km; or ), spelt kilometer in American and Philippine English, is a unit of length in the International System of Units (SI), equal to one thousand metres (kilo- being the SI prefix for ). It is the preferred measu ...
, in a matter of months. Burrowing behavior, in which an individual sinks itself entirely or partially into the substrate, is frequent among strombid gastropods. In the course of its life span, ''Lobatus galeatus'' spends part of the time partially buried in the sand.


Human uses

Giant conch shells were used as
wind instruments A wind instrument is a musical instrument that contains some type of resonator (usually a tube) in which a column of air is set into vibration by the player blowing into (or over) a mouthpiece set at or near the end of the resonator. The pitch ...
by the Chavín, a
pre-Incan This is a chart of cultural periods of Peru and the Andean Region developed by John Rowe and Edward Lanning and used by some archaeologists studying the area. An alternative dating system was developed by Luis Lumbreras and provides different ...
civilization that developed in the northern
Andean The Andes ( ), Andes Mountains or Andean Mountain Range (; ) are the longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range is long and wide (widest between 18°S ...
highlands of Peru. ''Lobatus galeatus'' shells were prepared for musical use through the creation of a mouthpiece. This was done by cutting a small tip of the
spire A spire is a tall, slender, pointed structure on top of a roof of a building or tower, especially at the summit of church steeples. A spire may have a square, circular, or polygonal plan, with a roughly conical or pyramidal shape. Spire ...
out and polishing the resulting hole, thus producing a trumpet-like instrument. In 2001, twenty such instruments were excavated from the Chavín de Huantar
archaeological site An archaeological site is a place (or group of physical sites) in which evidence of past activity is preserved (either prehistoric or recorded history, historic or contemporary), and which has been, or may be, investigated using the discipline ...
, which makes them nearly three thousand years old. This kind of playable shell is still used in the Andes, where they are known as ''pututu''. ''Lobatus galeatus'' is edible; it is an economically important species in many areas where it occurs. It is primarily used for subsistence and commercial fishery.


References

{{Taxonbar, from=Q3835936 Strombidae Gastropods described in 1823 Taxa named by William Swainson Peruvian musical instruments