Monstrilloida

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Monstrilloida
Monstrilla longiremis.jpg
Monstrilla longiremis
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Copepoda
Infraclass: Neocopepoda
Superorder: Podoplea
Order: Monstrilloida
G. O. Sars, 1901  [1]
Family: Monstrillidae
Dana, 1849  [2]

Monstrilloida is an order of copepods with a cosmopolitan distribution in the world's oceans. The order contains a single family, Monstrillidae. [1] The name of the first ever described genus Monstrilla is derived from latin, meaning "tiny monster", because the lack of usual diagnostic features of copepods puzzled early taxonomists. [3]

Contents

Description

The family Monstrillidae is characterised by having a well-developed fourth pair of swimming legs, but a rudimentary or absent fifth pair. Adults have no oral appendages, and the mouth leads only to a short, blind pharynx. They also lack second antennae, but show large, multiramous and setaceous antennulae. These antennulae are rigid and anteriorly oriented. Females carry a long pair of spines to which the eggs are attached, while males have a "genital protuberance, which is provided with lappets"; in both sexes, the genitalia are very different from those of all other copepods. [4] Some species have large, well-developed nauplius eyes, while others are basically blind.[ citation needed ]

Larval nauplial stages do not possess any discernible antennae, antenullae or mouth parts, but paired tube-shaped nourishing appendages to absorb nutrients from their host, which are also present in later copepodite stages that resemble the adult morphology; in adults, scars of these now discarded appendages remain as small processes on the cephalothorax. [5]

Taxonomy of genera and species descriptions are normally based on a few key characteristics: the length and setation-pattern of the antennulae, [6] presence/absence and morphology of the eyes, number and shape of caudal setae, and structure and setation-pattern of the fifth swimming leg in females/genital complex in males, respectively. [3]

Some taxonomically important features of Monstrillopsis planifrons, including armature of the right antennule Right antennule of Monstrillopsis planifrons.jpg
Some taxonomically important features of Monstrillopsis planifrons, including armature of the right antennule
Monstrilloid diversity: females of A. Cymbasoma cocoense B. Monstrillopsis chilensis C. Monstrillopsis igniterra D. Cymbasoma nicolettae ANT = antennules; AS = anal somite; CR =  caudal ramus; CS = caudal seta; EC = egg cluster; GS = genital somite; L1-L5 = legs 1-5; OC = ocellus; OP = oral papilla; OS = ovigerous spine; P2-P5 = pedigerous somites 2-5; VN = ventral nipple-like process. Monstrilloid diversity.png
Monstrilloid diversity: females of A. Cymbasoma cocoense B. Monstrillopsis chilensis C. Monstrillopsis igniterra D. Cymbasoma nicolettaeANT = antennules; AS = anal somite; CR =  caudal ramus; CS = caudal seta; EC = egg cluster; GS = genital somite; L1–L5 = legs 1–5; OC = ocellus; OP = oral papilla; OS = ovigerous spine; P2–P5 = pedigerous somites 2–5; VN = ventral nipple-like process.
Monstrilla sp., lateral view Monstrilla (YPM IZ 102076).jpeg
Monstrilla sp., lateral view


Distribution

Monstrilloids are distributed worldwide (including both the Arctic [7] and Antarctic), inhabiting coastal-neritic waters (0-200m depth). [8] Adults are regularly caught with plankton nets and are clearly pelagic organisms; however, the endoparasitic larval stages infect sessile benthic organisms and therefore are part of the epibenthos and hyperbenthos. [3]

Biology

Biologically and ecologically, our knowledge of the order is limited, although the life cycle differs from that of all other copepods: [9] Members of the Monstilloida are protelean parasites, meaning that their larval stages are parasitoids that kill their host to emerge as free-living subadults. Apparently, some hosts recover after the final subadult monstrilloid exits their body. [3] It is hypothesized, that the host's relative body size and the number and location of copepods parasitizing the same host determine whether it survives an infection. [3] The detailed life cycle may vary between different species, but generally follows a certain sequence: after a free-swimming, infective nauplius stage, the larvae develop inside benthic polychaetes, gastropods, sponges and bivalve mussels (They may be a pest in commercially important bivalve aquaculture), [10] from where the planktonic adults emerge. They do not possess functioning mouth-parts, their sole purpose is to reproduce. [11] In Contrast to holoplanktonic calanoid and cyclopoid copepods, Monstrilloids do not use their largest cephalic appendages, the antennulae, for locomotion, but to create a stream-line shaped corridor, rather using their four pairs of swimming legs to move in the water column. Sex determination depends on conspecifics infecting the same host individual. In case of 2-3 coexisting monstrilloids they become males, when there is only one parasite in a host, it develops into a female. [12]

different life cycle stages of Cymbasoma danae FMIB 47702 Stages in the life-history of Haemocera danae, one of the Monstrillidae.jpeg
different life cycle stages of Cymbasoma danae

Taxonomy

The taxonomy of the order and family is undergoing several revisions, [13] for instance, the family Thaumatopsyllidae was formerly included in the order, but is now usually placed in the Cyclopoida. [14] and the genus Strilloma is now considered a taxonomic synonym of Monstrilla , the largest genus. [15] In General, the Monstrilloida are taxonomically challenging, both regarding their relation to other copepod groups and species assignment within the order. Monstrilloida was placed as a sister taxa to the Siphonostomatoida, but a lack of mouth parts makes comparison based on homologies difficult. A more recent analysis placed the order nested within the fish-parasitizing caligiform groups of Siphonostomatoida. Consequently, they would have evolved from an ectoparasitic ancestor associated with fish; most parasitic copepods are not free-living as adults, so Monstrilloids presumably underwent a change in life cycle strategy, host selection and body morphology. [16] Yet the unique nature of the order Monstrilloida sister group of the Siphonostomatoida has been corroborated using modern molecular approaches. [17] So far, no ultimately satisfying copepod phylogeny has been proposed, and the placement of the monophyletic Monstrilloida remains unresolved. [5]

Because of their enigmatic life cycle, the morphology of most postnaupliar and copepodite stages is not known. Monstrilloids are not abundant in planktonic samples, and often only single specimens can be collected. Since many species occupy overlapping geographic ranges, males and females of different species collected in the same sample may be mistaken for conspecifics. Until now, both sexes were described for less than 25% of all known species. [5] To link males and females of a species, taxonomists have started to use molecular methods such as DNA Barcoding recently. [18]

As of 2019, the order Monstrilloida contains seven accepted genera with more than 160 species: [7] [19]

Related Research Articles

<span class="mw-page-title-main">Copepod</span> Subclass of crustaceans

Copepods are a group of small crustaceans found in nearly every freshwater and saltwater habitat. Some species are planktonic, some are benthic, a number of species have parasitic phases, and some continental species may live in limnoterrestrial habitats and other wet terrestrial places, such as swamps, under leaf fall in wet forests, bogs, springs, ephemeral ponds, puddles, damp moss, or water-filled recesses of plants (phytotelmata) such as bromeliads and pitcher plants. Many live underground in marine and freshwater caves, sinkholes, or stream beds. Copepods are sometimes used as biodiversity indicators.

<span class="mw-page-title-main">Tantulocarida</span> Subclass of crustaceans

Tantulocarida is a highly specialised group of parasitic crustaceans that consists of about 33 species, treated as a class in superclass Multicrustacea. They are typically ectoparasites that infest copepods, isopods, tanaids, amphipods and ostracods.

<span class="mw-page-title-main">Rhizocephala</span> Superorder of barnacles

Rhizocephala are derived barnacles that parasitise mostly decapod crustaceans, but can also infest Peracarida, mantis shrimps and thoracican barnacles, and are found from the deep ocean to freshwater. Together with their sister groups Thoracica and Acrothoracica, they make up the subclass Cirripedia. Their body plan is uniquely reduced in an extreme adaptation to their parasitic lifestyle, and makes their relationship to other barnacles unrecognisable in the adult form. The name Rhizocephala derives from the Ancient Greek roots ῥίζα and κεφαλή, describing the adult female, which mostly consists of a network of thread-like extensions penetrating the body of the host.

<span class="mw-page-title-main">Argulidae</span> Family of crustaceans

The family Argulidae, whose members are commonly known as carp lice or fish lice, are parasitic crustaceans in the class Ichthyostraca. It is the only family in the monotypic subclass Branchiura and the order Arguloida, although a second family, Dipteropeltidae, has been proposed. Although they are thought to be primitive forms, they have no fossil record.

<span class="mw-page-title-main">Poecilostomatoida</span> Suborder of crustaceans

Poecilostomatoida is a suborder of copepods. Although it was previously considered a separate order, recent research showed it to be nested within the Cyclopoida.

<i>Macrocyclops albidus</i> Species of crustacean

Macrocyclops albidus is a larvivorous copepod species.

Acartia hudsonica is a species of marine copepod belonging to the family Acartiidae. Acartia hudsonica is a coastal, cold water species that can be found along the northwest Atlantic coast.

<span class="mw-page-title-main">Crustacean larva</span> Crustacean larval and immature stages between hatching and adult form

Crustaceans may pass through a number of larval and immature stages between hatching from their eggs and reaching their adult form. Each of the stages is separated by a moult, in which the hard exoskeleton is shed to allow the animal to grow. The larvae of crustaceans often bear little resemblance to the adult, and there are still cases where it is not known what larvae will grow into what adults. This is especially true of crustaceans which live as benthic adults, more-so than where the larvae are planktonic, and thereby easily caught.

<span class="mw-page-title-main">Pennellidae</span> Family of crustaceans

Pennellidae is a family of parasitic copepods. When anchored on a host, they have a portion of the body on the outside of the host, whereas the remaining anterior part of the parasite is hidden inside tissues of the host.

<i>Lernaeocera branchialis</i> Species of crustacean

Lernaeocera branchialis, sometimes called cod worm, is a parasite of marine fish, found mainly in the North Atlantic. It is a marine copepod which starts life as a small pelagic crustacean larva. It is among the largest of copepods, ranging in size from 2 to 3 millimetres when it matures as a copepodid larva to more than 40 mm as a sessile adult.

<span class="mw-page-title-main">Crustacean</span> Subphylum of arthropods

Crustaceans belong to the subphylum Crustacea, and form a large, diverse group of arthropods including decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, opossum shrimps, amphipods and mantis shrimp. The crustacean group can be treated as a subphylum under the clade Mandibulata. It is now well accepted that the hexapods emerged deep in the Crustacean group, with the completed group referred to as Pancrustacea. The three classes Cephalocarida, Branchiopoda and Remipedia are more closely related to the hexapods than they are to any of the other crustaceans.

Peniculisa is a genus of marine parasitic copepods in the family Pennellidae.

<i>Protosarcotretes</i> Genus of crustaceans

Protosarcotretes is a genus of marine copepods in the family Pennellidae. Its type-species is Protosarcotretes nishikawai. This genus exhibits the most plesiomorphic states in the first to fourth legs of pennellids, and is differentiated from two closely related pennellid genera Sarcotretes and Lernaeenicus by the morphology of the oral appendages.

Blastodinium is a diverse genus of dinoflagellates and important parasites of planktonic copepods. They exist in either a parasitic stage, a trophont stage, and a dinospore stage. Although morphologically and functionally diverse, as parasites they live exclusively in the intestinal tract of copeods.

<i>Peniculus minuticaudae</i> Species of Maxillopoda

Peniculus minuticaudae is a species of parasitic pennellid copepod. It is known from the northeast Pacific Ocean. It was originally described in 1956, redescribed in 2012, and its complete life cycle has been elucidated on the cultured threadsail filefish, Stephanolepis cirrhifer in 2013.

<i>Peniculus hokutoae</i> Species of copepod

Peniculus hokutoae is a species of parasitic pennellid copepod. It was described in 2018 from a single female. The type-host is the myctophid fish Symbolophorus evermanni and the type-locality is off Japan. The Japanese name of this species is hokuto-kozutsu-hijikimushi.

Longipedia is a genus of marine copepods of the family Longipediidae, order Canuelloida.

Mesocyclops longisetus is a species of freshwater copepod in the family Cyclopidae. Two subspecies are accepted, Mesocyclops longisetus curvatus Dussart, 1987, and Mesocyclops longisetus longisetus. It has a neotropical distribution.

Ergasilus curticrus is a freshwater parasitic copepod named in 2015. Described from the Orinoco river basin, it was found solely to be hosted by individuals of the Characiform fish species Bryconops giacopinii. Of those located in South America, it is one of only five species in its genus to be found outside of Brazil.

References

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