''Lysiosquillina maculata'', the zebra mantis shrimp, striped mantis shrimp or razor mantis, is a species of

mantis shrimp Mantis shrimp, or stomatopods, are carnivorous marine crustaceans of the order Stomatopoda (). Stomatopods branched off from other members of the class Malacostraca around 340 million years ago. Mantis shrimp typically grow to around in length, ...

found across the

Indo-Pacific The Indo-Pacific is a vast biogeographic region of Earth. In a narrow sense, sometimes known as the Indo-West Pacific or Indo-Pacific Asia, it comprises the tropical waters of the Indian Ocean, the western and central Pacific Ocean, and the ...

region from

East Africa East Africa, Eastern Africa, or East of Africa, is the eastern subregion of the African continent. In the United Nations Statistics Division scheme of geographic regions, 10-11-(16*) territories make up Eastern Africa: Due to the historica ...

to the Galápagos and

Hawaiian Islands The Hawaiian Islands ( haw, Nā Mokupuni o Hawai‘i) are an archipelago of eight major islands, several atolls, and numerous smaller islets in the North Pacific Ocean, extending some from the island of Hawaii in the south to northernmost ...

. At a length up to 40 cm, ''L. maculata'' is the largest mantis shrimp in the world. ''L. maculata'' may be distinguished from its congener '' L. sulcata'' by the greater number of teeth on the last segment of its

raptorial The term ''raptorial'' implies much the same as ''predatory'' but most often refers to modifications of an arthropod's foreleg that make it function for the grasping of prey while it is consumed, where the gripping surfaces are formed from the o ...

claw A claw is a curved, pointed appendage found at the end of a toe or finger in most amniotes (mammals, reptiles, birds). Some invertebrates such as beetles and spiders have somewhat similar fine, hooked structures at the end of the leg or tarsus ...

, and by the colouration of the

uropod Uropods are posterior appendages found on a wide variety of crustaceans. They typically have functions in locomotion. Definition Uropods are often defined as the appendages of the last body segment of a crustacean. An alternative definition sugge ...

endopod The arthropod leg is a form of jointed appendage of arthropods, usually used for walking. Many of the terms used for arthropod leg segments (called podomeres) are of Latin origin, and may be confused with terms for bones: ''coxa'' (meaning hip (anat ...

, the distal half of which is dark in ''L. maculata'' but not in ''L. sulcata''. A small

artisan An artisan (from french: artisan, it, artigiano) is a skilled craft worker who makes or creates material objects partly or entirely by hand. These objects may be functional or strictly decorative, for example furniture, decorative art ...

fishery Fishery can mean either the enterprise of raising or harvesting fish and other aquatic life; or more commonly, the site where such enterprise takes place ( a.k.a. fishing ground). Commercial fisheries include wild fisheries and fish farms, ...

exists for this species.

Hunting strategies

Stomatopod Mantis shrimp, or stomatopods, are carnivorous marine crustaceans of the order Stomatopoda (). Stomatopods branched off from other members of the class Malacostraca around 340 million years ago. Mantis shrimp typically grow to around in length ...

s are distinguished by their unique hunting adaptations, the most obvious being their second maxilliped modified into a powerful raptorial claw. Whether a mantis shrimp is classified as a smasher or a spearer is distinguished based on the form of the raptorial claw and the way in which the mantis shrimp uses it. Spearers have unfolded dactyls that are lined with sharp spines and are especially elongated to reach prey. Smashers on the other hand have folded dactyls that are modified for smashing through their heavily calcified shell.Van Der Wal, C, et al. "The Evolutionary History of Stomatopoda (Crustacea: Malacostraca) Inferred from Molecular Data". ''Peerj'', vol. 5, no. 9, 2017, doi:10.7717/peerj.3844. The zebra mantis shrimp is an example of a "spearing" mantis shrimp.deVries, M. S., et al. "Strike Mechanics of an Ambush Predator: the Spearing Mantis Shrimp". ''Journal of Experimental Biology'', The Company of Biologists Ltd, 15 Dec. 2012, jeb.biologists.org/content/215/24/4374.

Predation

By hiding in sandy burrows, ''Lysiosquillina maculata'' strike soft-bodied evasive prey that swim by such as small fish and

malacostraca Malacostraca (from New Latin; ) is the largest of the six classes of crustaceans, containing about 40,000 living species, divided among 16 orders. Its members, the malacostracans, display a great diversity of body forms and include crabs, lobs ...

ns. In order to create high attack speeds, they have a spring and latch structure at the base of their raptorial appendages that allows them to create a spring-loaded strike. This strike system consists of a four-bar linkage mechanism and a coupled lever. By storing energy in this spring-like system a faster strike is created than would be possible from solely muscle action.Anderson, Philip S. L., et al. “LEVERS AND LINKAGES: MECHANICAL TRADE‐OFFS IN A POWER‐AMPLIFIED SYSTEM.” ''The Canadian Journal of Chemical Engineering'', Wiley-Blackwell, 5 May 2014, onlinelibrary.wiley.com/doi/full/10.1111/evo.12407. Prior to attacking prey, ''Lysiosquillina maculata'' only peeks its eyes above the entrance to the burrow. As soon as the prey move within striking distance of the burrows, the mantis shrimp lunges out and snares the prey with both of its raptorial appendages. Zebra mantis shrimp attack with a mean peak speed of 2.3 m/s and with a mean duration of 24.98 ms. This speed is significantly slower than those generated by the smashing mantis shrimp, whose strikes can reach 14–23 m/s. However, it is similar to those of other aquatic predators attacking evasive prey. This discrepancy is because spearing mantis shrimp display displacement amplification whereas smashing mantis shrimp display force amplification. This makes sense given their hunting strategies. “Smashers” need to apply a large amount of force, but easily get close to their sessile, hard-shelled prey. However, for “spearers” like the zebra mantis shrimp, it is more advantageous to have greater reach when targeting prey with their

ambush An ambush is a long-established military tactic in which a combatant uses an advantage of concealment or the element of surprise to attack unsuspecting enemy combatants from concealed positions, such as among dense underbrush or behind moun ...

attack strategy. In addition, it has been shown that larger mantis shrimp species strike more slowly, resulting in the slower speeds displayed by ''Lysiosquillina maculata''.

Eyesight

The class of

crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapoda, decapods, ostracoda, seed shrimp, branchiopoda, branchiopods, argulidae, fish lice, krill, remipedes, isopoda, isopods, barnacles, copepods, ...

s called

s, which includes ''Lysiosquillina maculata'' has the most complex visual receptors in the animal kingdom. Many species are thought to be able to express up to 16 different visual pigments. In addition,

s have a tripartite

cornea The cornea is the transparent front part of the eye that covers the iris, pupil, and anterior chamber. Along with the anterior chamber and lens, the cornea refracts light, accounting for approximately two-thirds of the eye's total optical ...

which contains upper and lower halves, separated by a middle band made up of

ommatidia The compound eyes of arthropods like insects, crustaceans and millipedes are composed of units called ommatidia (singular: ommatidium). An ommatidium contains a cluster of photoreceptor cells surrounded by support cells and pigment cells. The ...

. These photoreceptors include specialized receptors that can detect a wide range of visible and

ultraviolet light Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nm (with a corresponding frequency around 30 PHz) to 400 nm (750 THz), shorter than that of visible light, but longer than X-rays. UV radiatio ...

, as well as being able to detect linearly and circularly polarized light.

polarized light Polarization ( also polarisation) is a property applying to transverse waves that specifies the geometrical orientation of the oscillations. In a transverse wave, the direction of the oscillation is perpendicular to the direction of motion of t ...

is used by many insects for navigation, however, in other invertebrates like cephalopods and

s it is used primarily to increase visual contrast and for visual signaling.Templin, Rachel M., et al. “Fel.” Journal of Experimental Biology, The Company of Biologists Ltd, 15 Sept. 2017, jeb.biologists.org/content/220/18/3222.abstract. When searching for prey, each eye moves independently from the other. However, when a prey enters the visual field of one of the eyes, they lock into place focused on the prey.Dore, B., et al. “Growth and Light/Dark Adaptation in Lysiosquillina Maculata (Stomatopoda, Crustacea).” ''Journal of Biological Research'', vol. 81, no. 1, 2006, doi:10.4081/jbr.2006.8080.

Behavior

''Lysiosquillina maculata'' display social monogamy with maternal egg care, meaning the females are the ones taking care of the eggs. Social monogamy is usually attributed to the large and costly to construct burrows of these species. These burrows are used for protection, and as a place to hide and wait for prey. In addition, there is a large risk associated with finding mates. ''Lysiosquillina maculata'' also displays

sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most an ...

, with males having larger raptorial appendages, although males and females have similar overall body sizes.

Evolution

''Lysiosquillina maculata'' belongs to the

order Stomatopoda. Stomatopoda is made up of three distinct suborders, Palaeostomatopodea, Archaeostomatopodea, and Unipeltata. All living

stomatopod Mantis shrimp, or stomatopods, are carnivorous marine crustaceans of the order Stomatopoda (). Stomatopods branched off from other members of the class Malacostraca around 340 million years ago. Mantis shrimp typically grow to around in length ...

s belong to the suborder

Unipeltata Mantis shrimp, or stomatopods, are carnivorous marine crustaceans of the order Stomatopoda (). Stomatopods branched off from other members of the class Malacostraca around 340 million years ago. Mantis shrimp typically grow to around in length ...

which contains seven distinct superfamilies of mantis shrimp, made up of almost 500 species. Using molecular clocks, research has shown that around 340 million years ago,

s branched off from their closest

relatives. They also found that the specialized smashing appendage developed around 126 million years ago, after the spearing appendage which arose around 155 million years ago.

References

External links

* * {{Taxonbar, from=Q3855731 Stomatopoda Edible crustaceans Galápagos Islands coastal fauna Crustaceans described in 1793