Temporal range: Earliest Cambrian (Fortunian)–Recent
von Siebold, 1848
|Subphyla, unplaced genera, and classes|
|around 1,170,000 species.|
Condylipoda Latreille, 1802
Arthropods use combinations of
The evolutionary ancestry of arthropods dates back to the
The word arthropod comes from the
In common parlance, terrestrial arthropods are often called bugs.
Arthropods are invertebrates with segmented bodies and jointed limbs. The exoskeleton or cuticles consists of chitin, a polymer of N-Acetylglucosamine. The cuticle of many crustaceans, beetle mites, the clades Penetini and Archaeoglenini inside the beetle subfamily Phrenapatinae, and millipedes (except for bristly millipedes) is also biomineralized with calcium carbonate. Calcification of the endosternite, an internal structure used for muscle attachments, also occur in some opiliones, and the pupal cuticle of the fly Bactrocera dorsalis contains calcium phosphate.
Estimates of the number of arthropod species vary between 1,170,000 and 5 to 10 million and account for over 80 percent of all known living animal species. The number of species remains difficult to determine. This is due to the census modeling assumptions projected onto other regions in order to scale up from counts at specific locations applied to the whole world. A study in 1992 estimated that there were 500,000 species of animals and plants in Costa Rica alone, of which 365,000 were arthropods.
They are important members of marine, freshwater, land and air
The embryos of all arthropods are segmented, built from a series of repeated modules. The last common ancestor of living arthropods probably consisted of a series of undifferentiated segments, each with a pair of appendages that functioned as limbs. However, all known living and fossil arthropods have grouped segments into tagmata in which segments and their limbs are specialized in various ways.
The three-part appearance of many insect bodies and the two-part appearance of spiders is a result of this grouping. There are no external signs of segmentation in mites. Arthropods also have two body elements that are not part of this serially repeated pattern of segments, an ocular somite at the front, where the mouth and eyes originated, and a telson at the rear, behind the anus.
Originally it seems that each appendage-bearing segment had two separate pairs of appendages: an upper, unsegmented
The most conspicuous specialization of segments is in the head. The four major groups of arthropods –
Working out the evolutionary stages by which all these different combinations could have appeared is so difficult that it has long been known as "the arthropod head problem". In 1960, R. E. Snodgrass even hoped it would not be solved, as he found trying to work out solutions to be fun.[Note 2]
Arthropod exoskeletons are made of
The exoskeletons of most aquatic crustaceans are biomineralized with calcium carbonate extracted from the water. Some terrestrial crustaceans have developed means of storing the mineral, since on land they cannot rely on a steady supply of dissolved calcium carbonate. Biomineralization generally affects the exocuticle and the outer part of the endocuticle. Two recent hypotheses about the evolution of biomineralization in arthropods and other groups of animals propose that it provides tougher defensive armor, and that it allows animals to grow larger and stronger by providing more rigid skeletons; and in either case a mineral-organic composite exoskeleton is cheaper to build than an all-organic one of comparable strength.
The cuticle may have
Although all arthropods use muscles attached to the inside of the exoskeleton to flex their limbs, some still use
The exoskeleton cannot stretch and thus restricts growth. Arthropods, therefore, replace their exoskeletons by undergoing ecdysis (moulting), or shedding the old exoskeleton after growing a new one that is not yet hardened. Moulting cycles run nearly continuously until an arthropod reaches full size.
The developmental stages between each moult (ecdysis) until sexual maturity is reached is called an instar. Differences between instars can often be seen in altered body proportions, colors, patterns, changes in the number of body segments or head width. After moulting, i.e. shedding their exoskeleton, the juvenile arthropods continue in their life cycle until they either pupate or moult again.
In the initial phase of moulting, the animal stops feeding and its epidermis releases moulting fluid, a mixture of
Because arthropods are unprotected and nearly immobilized until the new cuticle has hardened, they are in danger both of being trapped in the old cuticle and of being attacked by predators. Moulting may be responsible for 80 to 90% of all arthropod deaths.
Arthropod bodies are also segmented internally, and the nervous, muscular, circulatory, and excretory systems have repeated components.
Respiration and circulation
Arthropods have open
The heart is typically a muscular tube that runs just under the back and for most of the length of the hemocoel. It contracts in ripples that run from rear to front, pushing blood forwards. Sections not being squeezed by the heart muscle are expanded either by elastic
Arthropods have a wide variety of respiratory systems. Small species often do not have any, since their high ratio of surface area to volume enables simple diffusion through the body surface to supply enough oxygen. Crustacea usually have gills that are modified appendages. Many arachnids have book lungs. Tracheae, systems of branching tunnels that run from the openings in the body walls, deliver oxygen directly to individual cells in many insects, myriapods and arachnids.
Living arthropods have paired main nerve cords running along their bodies below the gut, and in each segment the cords form a pair of
There are two different types of arthropod excretory systems. In aquatic arthropods, the end-product of biochemical reactions that
Most arthropods lack balance and
Most arthropods have sophisticated visual systems that include one or more usually both of
Compound eyes consist of fifteen to several thousand independent
Reproduction and development
A few arthropods, such as
Most arthropods lay eggs,
Last common ancestor
Based on the distribution of shared
It has been proposed that the Ediacaran animals Parvancorina and Spriggina, from around 555 million years ago, were arthropods, but later study shows that their affinities of being origin of arthropods are not reliable. Small arthropods with bivalve-like shells have been found in Early Cambrian fossil beds dating 541 to 539 million years ago in China and Australia. The earliest Cambrian trilobite fossils are about 530 million years old, but the class was already quite diverse and worldwide, suggesting that they had been around for quite some time. In the Maotianshan shales, which date to between 530 and 520 million years ago, fossils of arthropods such as Kylinxia and Erratus have been found that seem to show a transitional split between lobopodia and other more primitive stem arthropods. Re-examination in the 1970s of the Burgess Shale fossils from about 505 million years ago identified many arthropods, some of which could not be assigned to any of the well-known groups, and thus intensified the debate about the Cambrian explosion. A fossil of Marrella from the Burgess Shale has provided the earliest clear evidence of moulting.
The earliest fossil crustaceans date from about 511 million years ago in the Cambrian, and fossil shrimp from about 500 million years ago apparently formed a tight-knit procession across the seabed. Crustacean fossils are common from the Ordovician period onwards. They have remained almost entirely aquatic, possibly because they never developed excretory systems that conserve water. In 2020 scientists announced the discovery of Kylinxia, a five-eyed ~5 cm long shrimp-like animal living 518 Mya that – with multiple distinctive features – appears to be a key ‘missing link’ of the evolution from Anomalocaris to true arthropods and could be at the evolutionary root of true arthropods.
Arthropods provide the earliest identifiable fossils of land animals, from about 419 million years ago in the Late Silurian, and terrestrial tracks from about 450 million years ago appear to have been made by arthropods. Arthropods possessed attributes that were easy coopted for life on land; their existing jointed exoskeletons provided protection against desiccation, support against gravity and a means of locomotion that was not dependent on water. Around the same time the aquatic, scorpion-like eurypterids became the largest ever arthropods, some as long as 2.5 m (8 ft 2 in).
The oldest known
The oldest possible insect fossil is the
Evolutionary family tree
From 1952 to 1977, zoologist
Further analysis and discoveries in the 1990s reversed this view, and led to acceptance that arthropods are
A contrary view was presented in 2003, when Jan Bergström and Xian-Guang Hou argued that, if arthropods were a "sister-group" to any of the anomalocarids, they must have lost and then re-evolved features that were well-developed in the anomalocarids. The earliest known arthropods ate mud in order to extract food particles from it, and possessed variable numbers of segments with unspecialized appendages that functioned as both gills and legs. Anomalocarids were, by the standards of the time, huge and sophisticated predators with specialized mouths and grasping appendages, fixed numbers of segments some of which were specialized, tail fins, and gills that were very different from those of arthropods. In 2006, they suggested that arthropods were more closely related to
Higher up the "family tree", the
In the 1990s,
If the Ecdysozoa hypothesis is correct, then segmentation of arthropods and annelids either has evolved convergently or has been inherited from a much older ancestor and subsequently lost in several other lineages, such as the non-arthropod members of the Ecdysozoa.
Phylogeny of stem-group arthropods
Modern interpretations of the basal, extinct
- The "Giant" or "Siberiid Lobopodians", such as Jianshanopodia, Siberion and Megadictyon, are the most basal grade in the total-group Arthropoda.
- The "Gilled Lobopodians", such as Kerygmachela, Pambdelurion and Opabinia, are the second most basal grade.
- The Radiodonta, which traditionally known as anomalocaridids come in third position, and are thought to be monophyletic.
- A possible "upper stem-group" assemblage of more uncertain position but contained within Deuteropoda: the Fuxianhuiida, Megacheira, and multiple "bivalved forms" including Isoxyida (Isoxys and Surusicaris) and Hymenocarina.
However, recent analyses since late 2010s also show that these "upper stem-groups" might be inside the crown-group:
The following cladogram shows the probable relationships between crown-group Arthropoda and stem-group Arthropoda according to O’Flynn et al. 2022, including two new fossils found to be the most early branches of Deuteropoda (the "upper stem-groups" in previous studies are marked in asterisk, living groups are marked in bold):
Note that the subphylum
Phylogeny of living arthropods
The following cladogram shows the internal relationships between all the living classes of arthropods as of late 2010s, as well as the estimated timing for some of the clades:
The phylum Arthropoda is typically
- harvestmen, spiders, scorpions and related organisms characterized by the presence of chelicerae, appendages just above/in front of the mouthparts. Chelicerae appear in scorpions and horseshoe crabs as tiny claws that they use in feeding, but those of spiders have developed as fangs that inject venom.
- body segmentseach of which bearing one or two pairs of legs (or in a few cases being legless). All members are exclusively terrestrial.
Aside from these major groups, a number of fossil forms, mostly from the early Cambrian period, are difficult to place taxonomically, either from lack of obvious affinity to any of the main groups or from clear affinity to several of them. Marrella was the first one to be recognized as significantly different from the well-known groups.
The phylogeny of the major extant arthropod groups has been an area of considerable interest and dispute. Recent studies strongly suggest that Crustacea, as traditionally defined, is paraphyletic, with Hexapoda having evolved from within it, so that Crustacea and Hexapoda form a clade, Pancrustacea. The position of Myriapoda, Chelicerata and Pancrustacea remains unclear as of April 2012[update]. In some studies, Myriapoda is grouped with Chelicerata (forming Myriochelata); in other studies, Myriapoda is grouped with Pancrustacea (forming Mandibulata), or Myriapoda may be sister to Chelicerata plus Pancrustacea.
The placement of the extinct trilobites is also a frequent subject of dispute. One of the newer hypotheses is that the chelicerae have originated from the same pair of appendages that evolved into antennae in the ancestors of Mandibulata, which would place trilobites, which had antennae, closer to Mandibulata than Chelicerata.
Interaction with humans
However, the greatest contribution of arthropods to human food supply is by pollination: a 2008 study examined the 100 crops that FAO lists as grown for food, and estimated pollination's economic value as €153 billion, or 9.5 per cent of the value of world agricultural production used for human food in 2005. Besides pollinating, bees produce honey, which is the basis of a rapidly growing industry and international trade.
The red dye
The relative simplicity of the arthropods' body plan, allowing them to move on a variety of surfaces both on land and in water, have made them useful as models for
|Disease||Insect||Cases per year||Deaths per year|
|Malaria||Anopheles mosquito||267 M||1 to 2 M|
|Dengue fever||Aedes mosquito||?||?|
|Yellow fever||Aedes mosquito||4,432||1,177|
|Filariasis||Culex mosquito||250 M||unknown|
Although arthropods are the most numerous phylum on Earth, and thousands of arthropod species are venomous, they inflict relatively few serious bites and stings on humans. Far more serious are the effects on humans of diseases like
Many species of arthropods, principally insects but also mites, are agricultural and forest pests.
Even amongst arthropods usually thought of as obligate
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