Annotated classification

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Also known as: Crustacea

The following classification is based largely on that given in Synopsis and Classification of Living Organisms (1982) but has been modified to take account of advances made since that date. Groups marked with a dagger (†) are extinct and known only from fossils.

  • Subphylum Crustacea
    Two pairs of sensory appendages in front of mouth, and 3 pairs of jaws behind mouth; some parasitic and lack all appendages when adult; mostly aquatic; about 45,000 species known.
    • Class Cephalocarida (horseshoe shrimps)
      Holocene; primitive; blind; head shield without carapace; maxilla and all trunk limbs alike, with jointed inner branch and leaflike outer branches; abdominal segments without limbs; telson and furca present; length about 3 mm; marine, intertidal down to 300 m; only 9 known species.
    • Class Branchiopoda
      Early Devonian to present; limbs usually leaflike; maxillae reduced; eyes sometimes stalked, usually sessile (unstalked), often fused to form a single large median eye; nauplius, but some with direct development; predominantly freshwater, some marine, and some in strong inland brines; about 900 species.
    • Class Remipedia
      Holocene; body elongated; more than 30 segments, each with biramous appendages projecting sideways; antennules biramous; maxillules, maxillae, and maxillipeds uniramous and grasping; marine cave dwellers; about 17 species.
      • †Order Enantiopoda
        Carboniferous; single fossil, Tesnusocaris.
    • Class Maxillopoda
      Five pairs of head appendages; single, simple, median eye; antennules uniramous; maxillae usually present; up to 11 trunk segments; over 23,000 species.
      • Subclass Thecostraca
        Bivalved carapace of cypris larva forms an enveloping mantle in the adult; parasitic forms recognizable only by larval stages.
      • Subclass Cirripedia (barnacles)
        Late Silurian to present; sedentary; 6 pairs of trunk limbs (cirri); larvae free-swimming; sessile adults with carapace developed into a mantle; about 1,100 species.
        • Order Ascothoracica
          Cretaceous to present; parasites on sea anemones and echinoderms; body typically enclosed in a bivalved carapace; some with segmented abdomen and caudal furca; others distorted by outgrowths of the gut and ovary, giving a bushlike appearance; males dwarfed, living in mantle cavities of females; marine; about 30 species.
        • Order Rhizocephala
          Parasites on other crustaceans, mostly decapods; larvae typical nauplii and cyprids; adults ramify inside hosts and produce 1 or more reproductive bodies outside the host; marine; about 230 species.
        • Order Thoracica
          Silurian to present; the true barnacles; most are nonparasitic; larvae are nauplii and cyprids; adult body typically contained within calcareous shell plates; about 800 species.
      • Subclass Tantulocarida
        Holocene; eggs give rise to a tantulus larva with head shield and 6 pairs of thoracic limbs; adult females form large dorsal trunk sac between head shield and trunk, often losing the trunk; males with 6 pairs of trunk limbs; parasites on other crustaceans; marine; about 10 species.
      • Subclass Branchiura
        All species are ectoparasites on freshwater or marine fish; 125 species.
        • Order Arguloida (fish lice)
          Wide, flat carapace; paired compound eyes; unsegmented abdomen; 4 pairs of trunk limbs; fish parasites; capable of free swimming; mostly freshwater but some marine; about 125 species.
      • †Subclass Skaracarida
        Late Cambrian; 12 trunk segments; no thoracic appendages apart from maxillipeds.
      • Subclass Copepoda
        Miocene to present; no carapace; no compound eyes; 1 or more trunk segments fused to head; typically 6 pairs of thoracic limbs; no abdominal limbs; larva usually a nauplius; free-living and parasitic; worldwide; marine, freshwater, and some semi-terrestrial; at least 8,500 species.
        • Order Calanoida
          Antennules long, usually held stiffly at right angles to the length of the body; heart present; thorax articulates with a much narrower abdomen; fifth leg biramous; worldwide; marine and freshwater; mostly planktonic; about 2,000 species.
        • Order Misophrioida
          Carapace-like extension from the head covers the first segment bearing a swimming leg; heart present in some; no eyes; antennule with up to 27 segments; fifth leg biramous; marine.
        • Order Mormonilloida
          Antennule with 3 or 4 long segments and long setae; fifth leg absent; marine.
        • Order Harpacticoida
          Antennules short; abdomen not markedly narrower than the thorax; articulation between thoracic segments 5 and 6; mostly benthic, some tunnel in the fronds of seaweeds; usually 1 egg sac but some with 2; marine and freshwater, with some semiterrestrial on damp forest floors; about 2,250 species.
        • Order Cyclopoida
          Antennules medium length; thorax wider than abdomen; articulation between thoracic segments 5 and 6; mandibles with biting or chewing processes; eggs normally carried in 2 egg sacs; fifth leg uniramous; marine and freshwater; more than 3,000 species.
        • Order Poecilostomatoida
          Parasites and commensals of fish and invertebrates; mouth not tubelike or suckerlike; mandibles reduced; adult segmentation often reduced or lost; mostly marine, few freshwater.
        • Order Siphonostomatoida
          Mouth tubelike or forms a sucker with styletlike mandibles; adult segmentation reduced or lost; parasites and commensals on fish and invertebrates; mostly marine, some freshwater.
        • Order Monstrilloida
          Parasites on marine worms and mollusks; adults free-swimming; lack mouthparts and gut; biramous swimming legs; about 80 species.
      • Subclass Mystacocarida (mustache shrimps)
        Elongated; blind forms living in spaces between sand grains; antennules uniramous; antennae and mandibles biramous with long branches extending sideways; trunk limbs vestigial but caudal rami well-developed and pincerlike; marine; about 9 species.
    • Class Ostracoda (mussel or seed shrimps)
      Cambrian to present; body short; bivalved carapace encloses trunk and limbs; living forms have up to 7 pairs of appendages; most fossils known only from shells (carapaces); marine, freshwater, and some terrestrial; more than 2,000 living species worldwide.
      • †Order Bradoriida
        Cambrian to Ordovician.
      • †Order Phosphatocopida
        Cambrian; remarkable fossils with up to 9 pairs of well-preserved appendages.
      • †Order Leperditicopida
        Cambrian to Devonian.
      • †Order Beyrichicopida
        Silurian to Carboniferous.
      • Subclass Myodocopa
        • Order Myodocopida
          Silurian to present; antennal notch in shell; 5 pairs of postoral appendages; maxilla with a large respiratory plate; eyes usually present; marine.
        • Order Halocyprida
          Silurian to present; 5 pairs of postoral appendages; maxilla leglike; no eyes; marine.
          • Suborder Cladocopina
            Silurian to present; only 3 pairs of postoral appendages; marine.
      • Subclass Podocopa
        • Order Platycopida
          Ordovician to present; antennae biramous; 4 pairs of postoral limbs; marine.
        • Order Podocopida
          Ordovician to present; antennae uniramous; 5 pairs of postoral appendages; marine, freshwater, and terrestrial.
    • Class Malacostraca
      Cambrian to present; typically with compound eyes, stalked or sessile; 8 thoracic and 6 abdominal segments, each potentially capable of bearing a pair of appendages; about 22,000 species.
      • Subclass Phyllocarida
        Early Cambrian to present.
        • †Order Archaeostraca
          Devonian to Triassic.
        • †Order Hoplostraca
          Carboniferous.
        • Order Leptostraca
          Permian to present; bivalved carapace encloses 8 pairs of leaflike limbs; movable rostrum; telson with caudal rami; marine; about 10 species.
      • Subclass Hoplocarida
        Carboniferous to present.
        • Order Stomatopoda (mantis shrimps)
          Jurassic to present; eyes stalked; 2 movable segments in head; carapace leaves 4 thoracic segments uncovered; second thoracic limbs massive; marine; about 350 species.
        • †Order Palaeostomatopoda
          Carboniferous.
        • †Order Aeschronectida
          Carboniferous.
      • Subclass Eumalacostraca
        Late Devonian to Holocene; carapace (when present) not bivalved; rostrum fixed; first antenna 2-branched; thoracic legs with slender, many-segmented outer branch and stout, 7-segmented inner branch, often pincerlike, used in walking or food-gathering; 6 (rarely 7) abdominal segments, with pleopods and terminal uropods.
        • Superorder Syncarida
          Carboniferous to present; no carapace.
          • †Order Palaeocaridacea
            Carboniferous to Permian; first thoracic segment not fused to head; abdominal pleopods 2-branched, flaplike; 4 families.
          • Order Anaspidacea
            Permian to present; with or without eyes; antennules biramous; abdominal appendages well-developed; telson without a furca; South Australia and Tasmania; freshwater; about 8 species.
          • Order Stygocaridacea
            Blind, elongated forms with a small rostrum; first thoracic segment fused to head but sixth abdominal segment free; furca present; abdominal appendages reduced or absent; South America and New Zealand; freshwater, in spaces between sand grains; about 5 species.
          • Order Bathynellacea
            Blind, elongated forms, without a rostrum; first thoracic segment not fused to head but sixth abdominal segment fused with telson; antennules uniramous; worldwide; freshwater, in spaces between sand grains; about 100 species.
        • Superorder Peracarida
          Females with a ventral brood pouch formed by plates at the bases of some of the thoracic limbs; development direct, with offspring resembling adults.
          • Order Mysidacea (opossum shrimps)
            Triassic to present; carapace well-developed, covering most of thorax; 3–30 mm, with a few much larger; worldwide; mainly marine but some in brackish and fresh water; about 780 species.
          • Order Cumacea
            Permian to present; head and carapace much wider than trunk; uropods long and rodlike; 1–35 mm; marine; about 800 species.
          • Order Spelaeogriphacea
            Holocene; carapace short, fused to first and covering part of second thoracic segment; 4 pairs of well-developed abdominal appendages; about 8 mm; cave-dwelling; South Africa; freshwater; 1 species.
          • Order Mictacea
            Holocene; no functional eyes; carapace forms small lateral folds covering bases of mouthparts and maxillipeds; all trunk segments free; antennules biramous; thoracic limbs with exopods; abdominal appendages reduced, uniramous; 2.7–3.5 mm; deep-sea or in marine caves; 2 species.
          • Order Tanaidacea
            Permian to present; carapace short, fused to first 2 thoracic segments; second pair of thoracic limbs usually with pincers; abdomen short, usually with 5 pairs of biramous appendages; 2–25 mm; mainly marine; about 500 species.
          • Order Isopoda (pill bugs, wood lice, sea slaters)
            Carboniferous to present; eyes sessile; no carapace; abdominal appendages flattened and respiratory; thoracic limbs without exopods; some parasites highly modified as adults; most species 5–30 mm but some up to 270 mm; worldwide; marine, freshwater, and terrestrial; about 4,000 species.
          • Order Amphipoda (beach hoppers, scuds, well shrimps)
            Eocene to present; eyes sessile; no carapace; thoracic limbs have respiratory plates at base; few parasites; most 5–50 mm but up to 140 mm; worldwide; mainly marine but also numerous in fresh water; about 6,000 species.
        • Superorder Eucarida.
          Carapace large, fused dorsally to all thoracic segments; eyes stalked; development usually involves larval forms but is sometimes direct.
          • Order Euphausiacea (krill)
            Holocene; carapace does not cover gills; thoracic limbs with 2 well-developed branches; eggs usually shed freely; first larva a nauplius; 6–81 mm; worldwide; marine; about 85 species.
          • Order Amphionidacea
            Holocene; carapace large; mandible and maxillule vestigial; thoracic limbs with small outer branch; ventral brood pouch formed by large forwardly projecting first abdominal appendages; 2–3 cm; worldwide; marine, pelagic; 1 species.
          • Order Decapoda (shrimps, prawns, lobsters, crayfish, crabs)
            Devonian to present; carapace large, enclosing gills; first 3 pairs of thoracic appendages modified for feeding (maxillipeds); eggs often attached to abdominal appendages; worldwide; mostly marine but also freshwater and a few terrestrial; about 10,000 species.
        • Superorder Pancarida
          • Order Thermosbaenacea
            Holocene; eyes reduced or absent; brood pouch formed from dorsal extension of carapace; length about 4 mm; fresh and brackish water, some in warm springs; about 9 species.

Critical appraisal

There is no universal agreement on the classification of the Crustacea and even less agreement on the interrelationships between the various groups. Alternative classifications of the classes Branchiopoda and Malacostraca are discussed below. Some authorities, such as the author of the Cirripedes below, rank the cirripedes as a subclass. There is also some disagreement about the limits of the class Maxillopoda. Some would include the class Cephalocarida, others would exclude the class Ostracoda, and yet others do not regard the Maxillopoda as a valid group and would raise the maxilloped subclasses Copepoda and Ostracoda to separate classes. Some of the parasitic forms are sometimes separated and ranked as separate orders.