Boleophthalmus Pectinirostris
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''Boleophthalmus pectinirostris'', commonly known as the great blue spotted mudskipper, is a species of
mudskipper Mudskippers are any of the 23 extant species of amphibious fish from the subfamily Oxudercinae of the goby family (biology), family Oxudercidae. They are known for their unusual body shapes, preferences for semiaquatic habitats, limited terrestria ...
native to the north-western Pacific Ocean. It can be found on the coastlines of
Japan Japan is an island country in East Asia. Located in the Pacific Ocean off the northeast coast of the Asia, Asian mainland, it is bordered on the west by the Sea of Japan and extends from the Sea of Okhotsk in the north to the East China Sea ...
, eastern
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
,
Sumatra Sumatra () is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the list of islands by area, sixth-largest island in the world at 482,286.55 km2 (182,812 mi. ...
,
Malaysia Malaysia is a country in Southeast Asia. Featuring the Tanjung Piai, southernmost point of continental Eurasia, it is a federation, federal constitutional monarchy consisting of States and federal territories of Malaysia, 13 states and thre ...
,
Taiwan Taiwan, officially the Republic of China (ROC), is a country in East Asia. The main geography of Taiwan, island of Taiwan, also known as ''Formosa'', lies between the East China Sea, East and South China Seas in the northwestern Pacific Ocea ...
and the
Korean Peninsula Korea is a peninsular region in East Asia consisting of the Korean Peninsula, Jeju Island, and smaller islands. Since the end of World War II in 1945, it has been politically divided at or near the 38th parallel between North Korea (Dem ...
.


Description

Individuals of ''B. pectinirostris'' are predominantly greenish-grey in colour, with prominent sky blue speckles across their body, including their fins and on the skin below their eyes. They can grow all the way up to .


Anatomy


Epidermis

''B. pectinirostis'' is amphibious, and breathes through its
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and Subcutaneous tissue, hypodermis. The epidermal layer provides a barrier to infection from environmental pathogens and regulates the ...
, a process known as
cutaneous respiration Cutaneous respiration, or cutaneous gas exchange (sometimes called skin breathing), is a form of respiration in which gas exchange occurs across the skin or outer integument of an organism rather than gills or lungs. Cutaneous respiration may be ...
. A study examining specimens from
Jeollanam-do South Jeolla Province (), formerly South Chŏlla Province, also known as Jeonnam (), is a province in the Honam region, South Korea, and the southernmost province in mainland Korea. South Jeolla borders the provinces of North Jeolla to the nor ...
, South Korea, found that the epidermis of ''B. pectinirostis'' has three layers. The outermost layer consists of flattened, polygonal cells in one to eight layers of cells. The small cells are 5–7 μm by 5–8 μm, arranged in regular, compact rows on the outermost region of the layer, mostly in the jaw and fin epidermis. Larger
mucous Mucus (, ) is a slippery aqueous secretion produced by, and covering, mucous membranes. It is typically produced from cells found in mucous glands, although it may also originate from mixed glands, which contain both serous and mucous cells. It ...
cells are distributed throughout all regions of the epidermis except the fins and the sucking disc, located in between dermal bulges. The middle layer consists of layers of small cells and larger cells known as swollen cells that have a large
vacuole A vacuole () is a membrane-bound organelle which is present in Plant cell, plant and Fungus, fungal Cell (biology), cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water ...
. Whereas other air-breathing fishes have thicker middle layers with cells that uptake oxygen, members of the ''Boleophthalamus'' genus have the simple swollen cell structure that prevents dehydration while walking on land. The number of layers ranged from 1 to 15. The innermost layer is the stratus germinativum, and consists of one layer of
cuboidal Epithelium or epithelial tissue is a thin, continuous, protective layer of cells with little extracellular matrix. An example is the epidermis, the outermost layer of the skin. Epithelial ( mesothelial) tissues line the outer surfaces of many ...
and roughly
columnar Epithelium or epithelial tissue is a thin, continuous, protective layer of cells with little extracellular matrix. An example is the epidermis, the outermost layer of the skin. Epithelial ( mesothelial) tissues line the outer surfaces of many ...
basal cells The stratum basale (basal layer, sometimes referred to as ''stratum germinativum'') is the deepest layer of the five layers of the epidermis, the external covering of skin in mammals. The stratum basale is a single layer of columnar or cuboida ...
. The thickness of the epidermis largely depends on the middle layer thickness, and is generally thick in the body and thin in the fins.


Dermal bulges

''B. pectinirostis'' also has dermal bulges at the highest point of its scales on its body, except for where the fins and sucking disc are. The bulges' mean height ranges from 82 to 391 μm and their mean weight ranges from 172 to 485 μm. The distribution of bulges ranges from 0 to 6 per millimetre. The epidermis is very thin at these bulges, such that the stratus germinativum is very close to the outside.


Blood vessels

''B. pectinirostis'' has
blood vessels Blood vessels are the tubular structures of a circulatory system that transport blood throughout many animals’ bodies. Blood vessels transport blood cells, nutrients, and oxygen to most of the tissues of a body. They also take waste an ...
and dermal
capillaries A capillary is a small blood vessel, from 5 to 10 micrometres in diameter, and is part of the microcirculation system. Capillaries are microvessels and the smallest blood vessels in the body. They are composed of only the tunica intima (the in ...
in its dermis bulges just below the inner stratus germinativum layer.


Taxonomy


Etymology

''B. pectinirostriss generic name, ''Boleophthalmus'', is derived from the placement of the fish's eyes, which can be raised above the level of their
orbit In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an ...
s. It is taken from the Greek ''bole'' ('ejected') and ''ophthalmon'' ('eye'). The specific epithet is a compound
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
form, taken from ''pecten'' ('comb, rake') and ''rostrum'' ('beak'), and likely refers to the incised teeth of the fish's mandible, which it uses to scrape its food off the surface of the ground when on land.


Phylogeny

Phylogenetic analysis 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 data ...
of ''B. pectinirostris'' conducted in 2014 found that the species may in fact include at least two
cryptic species In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
as part of a
species complex In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
; one being native to the
South South is one of the cardinal directions or compass points. The direction is the opposite of north and is perpendicular to both west and east. Etymology The word ''south'' comes from Old English ''sūþ'', from earlier Proto-Germanic ''*sunþa ...
and
East China Sea The East China Sea is a marginal sea of the Western Pacific Ocean, located directly offshore from East China. China names the body of water along its eastern coast as "East Sea" (, ) due to direction, the name of "East China Sea" is otherwise ...
, and another being found in the
Strait of Malacca The Strait of Malacca is a narrow stretch of water, long and from wide, between the Malay Peninsula to the northeast and the Indonesian island of Sumatra to the southwest, connecting the Andaman Sea (Indian Ocean) and the South China Sea (Pa ...
northward towards Taiwan.


Habitat and distribution

''Boleophthalmus pectinirostris'' can be found in
Peninsular A peninsula is a landform that extends from a mainland and is only connected to land on one side. Peninsulas exist on each continent. The largest peninsula in the world is the Arabian Peninsula. Etymology The word ''peninsula'' derives , . ...
and eastern Malaysia, Sumatra (Indonesia), China, Taiwan, Korea, and Japan, living in warm
tropical The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
and
sub-tropical The subtropical zones or subtropics are geographical and climate zones immediately to the north and south of the tropics. Geographically part of the temperate zones of both hemispheres, they cover the middle latitudes from to approximately 3 ...
estuaries An estuary is a partially enclosed coastal body of brackish water with one or more rivers or streams flowing into it, and with a free connection to the open sea. Estuaries form a transition zone between river environments and maritime environm ...
. As a
euryhaline Euryhaline organisms are able to adapt to a wide range of salinities. An example of a euryhaline fish is the short-finned molly, '' Poecilia sphenops'', which can live in fresh water, brackish water, or salt water. The green crab ('' Carcinus m ...
species, it can also survive in a wide range of salinities. It prefers muddy areas near river mouths and
mangroves A mangrove is a shrub or tree that grows mainly in coastal 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 and remove sal ...
, where it is easier to burrow in. Its presence in Malaysia and Sumatra was confirmed in 2009, where it is
sympatric In biology, two closely related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter each other. An initially interbreeding population that splits into two or more distinct spe ...
with ''
Boleophthalmus boddarti ''Boleophthalmus boddarti'', commonly known as Boddart's goggle-eyed goby, is a species of mudskipper native to the Indo-Pacific, and the type species of the genus ''Boleophthalmus''. In Vietnam, this species is called "Cá Bống Sao", which me ...
''. Prior to 2009, the populations in this region were misidentified as the species ''
Boleophthalmus dussumieri ''Boleophthalmus dussumieri'', Dussumier's mudskipper, is a species of mudskipper native to the Indian Ocean where it can be found on mudflats in fresh, brackish and marine waters of Iraq, Pakistan and India as well as probably in Bangladesh. T ...
'', another species in the same genus which is related to both ''B. boddarti'' and ''B. pectiniriostris''.


Behaviour


Burrowing

Like other mudskippers, ''B. pectinirostris'' is able to move on land, using its pectoral and pelvic fins to clamber over the surface of
tidal flats Mudflats or mud flats, also known as tidal flats or, in Ireland, slob or slobs, are coastal wetlands that form in intertidal areas where sediments have been deposited by tides or rivers. A global analysis published in 2019 suggested that tidal ...
in its habitat at low tide. Using its tail, it can also hop across the ground to move faster. During high tide, the night, or when its predators are present, the fish retreats into a burrow, dug to a depth of approximately below the surface, becoming active again during the day or when the threat has passed. ''Boleophthalmus pectinirostris'' use their mouths to move soil and maintain their burrows, in the process increasing sediment surface area and oxygenating deeper layers of sediment. Burrows of the ''B. pectinirostris'' examined in
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 ...
ecosystems in Pandansari Brebes,
Central Java Central Java (, ) is a Provinces of Indonesia, province of Indonesia, located in the middle of the island of Java. Its administrative capital is Semarang. It is bordered by West Java in the west, the Indian Ocean and the Special Region of Yogya ...
were observed to have diameters of in muddy beach areas and in mangrove areas, identified as belonging to members of the species by tracks left by their fins. ''Boleophthalmus pectinirostris'' in Funing Bay,
Fujian, China Fujian is a province in southeastern China. Fujian is bordered by Zhejiang to the north, Jiangxi to the west, Guangdong to the south, and the Taiwan Strait to the east. Its capital is Fuzhou and its largest prefecture city by population is Qua ...
also constructs mud walls around the entrance of their burrows in the winter, creating a shallow walled pool that maintains a relatively consistent temperature, maintains a microphytobenthos (e.g.
diatoms A diatom (Neo-Latin ''diatoma'') is any member of a large group comprising several Genus, genera of algae, specifically microalgae, found in the oceans, waterways and soils of the world. Living diatoms make up a significant portion of Earth's B ...
) population for food, keeps other fish out, and prevents tides from moving the sediment around and in their burrows.


Territorialism

Great blue mudskippers are territorial, and males will fight with others of their species over access to burrows and during the breeding season, signalling their aggression by raising their large dorsal fins. The species is also known to compete with the Japanese mud crab (''Macrophthalmus japonicus'') over food in locations where both species occur.


Diet

''Boleophthalmus pectinirostris'' are
herbivore 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 ...
s. During low tide, they leave their tunnels to graze on diatoms at the surface.


Relationship with humans


Culinary use

''Boleophthalmus pectinirostris'' is of culinary importance in Japan, and is fished in areas where the species is particularly abundant, such as the
Ariake Sea The is a body of salt water surrounded by Fukuoka, Saga, Nagasaki, and Kumamoto Prefectures, all of which lie on the island of Kyūshū in Japan. It is the largest bay in Kyūshū. Its deepest point is only about 50 meters (165 ft) deep, and ext ...
and the
Yatsushiro Sea The , also called the , is a shallow semi-enclosed inland sea separating the island of Kyūshū from the Amakusa Islands. It lies mostly within Kumamoto Prefecture and at the southern end of the sea it also borders Kagoshima Prefecture. To the n ...
in
Saga Prefecture is a Prefectures of Japan, prefecture of Japan located on the island of Kyushu. Saga Prefecture has a population of roughly 780,000 and has a geographic area of . Saga Prefecture borders Fukuoka Prefecture to the northeast and Nagasaki Prefect ...
and
Kumamoto Prefecture is a Prefectures of Japan, prefecture of Japan located on the island of Kyūshū. Kumamoto Prefecture has a population of 1,748,134 () and has a geographic area of . Kumamoto Prefecture borders Fukuoka Prefecture to the north, Ōita Prefecture t ...
on the island of
Kyushu is the third-largest island of Japan's Japanese archipelago, four main islands and the most southerly of the four largest islands (i.e. excluding Okinawa Island, Okinawa and the other Ryukyu Islands, Ryukyu (''Nansei'') Ryukyu Islands, Islands ...
. A style of cooking the fish local to this region involves
grilling Grilling is a form of cooking that involves heat applied to the surface of food, commonly from above, below or from the side. Grilling usually involves a significant amount of direct, radiant heat, and tends to be used for cooking meat and v ...
and basting it whole over
charcoal Charcoal is a lightweight black carbon residue produced by strongly heating wood (or other animal and plant materials) in minimal oxygen to remove all water and volatile constituents. In the traditional version of this pyrolysis process, ca ...
.


Fishing

''Boleophthalmus pectinirostris'' is typically caught by line or by trapping. The line method, referred to as ''mutsukake'', is an unusual method developed in Japan that bears a resemblance to
fly fishing Fly fishing is an angling technique that uses an ultra-lightweight lure called an artificial fly, which typically mimics small invertebrates such as flying and aquatic insects to attract and catch fish. Because the mass of the fly lure is in ...
, and is carried out at low tide with the use of a long fishing rod, a harpoon-like hook that catches in the fish's skin, and a wooden
sled A sled, skid, sledge, or sleigh is a land vehicle that slides across a surface, usually of ice or snow. It is built with either a smooth underside or a separate body supported by two or more smooth, relatively narrow, longitudinal runners ...
called an ''oshiita'', which is used to travel out onto the tidal flats and to support oneself, as the surface of the flats is too soft to stand on. An alternative means of catching great blue spotted mudskippers is by using long burrow traps made out of hollow bamboo, which are driven into the tidal flat and trap the fish after they enter.


Status and conservation

As a shoreline species, ''B. pectinirostris'' is vulnerable to
water pollution Water pollution (or aquatic pollution) is the contamination of Body of water, water bodies, with a negative impact on their uses. It is usually a result of human activities. Water bodies include lakes, rivers, oceans, aquifers, reservoirs and ...
and
overfishing Overfishing is the removal of a species of fish (i.e. fishing) from a body of water at a rate greater than that the species can replenish its population naturally (i.e. the overexploitation of the fishery's existing Fish stocks, fish stock), resu ...
, and had formerly been in decline from around 1970 due to these factors. However, surveys of the populations in the Ariake and Yatsushiro Sea conducted in 2013 found that the species' numbers had increased by over eight times since the previous survey, which was undertaken 16 years prior. ''B. pectinirostris'''s recovery is believed to be the result of improved water quality, reduced fishing of the species due to a lower demand, and implementation of conservation measures. A series of mild winters may have also factored into the population increase.


References

{{Taxonbar, from=Q1186232 pectinirostris Mudskippers Fish of East Asia Fish of Japan Fish of the Pacific Ocean Taxa named by Carl Linnaeus Fish described in 1758