Arthropods: Habitat, Classification and Phylogeny
Arthropods: Habitat, Classification and Phylogeny
Phylogeny
Article Shared by <="" div="">
ADVERTISEMENTS:
In this article we will discuss about Arthropods:- 1. Taxonomic History of Arthropods 2.
Definition of Arthropods 3. Habit and Habitat 4. Classification 5. Phylogeny.
Taxonomic History of Arthropods:
The earliest record of the study of arthropods is available from the work of Aristotle (384-322
B.C.), who coined the term Malacostraca to include crabs and the related forms.
The present trend of studying Arthropoda began with the work of Linnaeus (1707-1778), who
created a group Insecta aptera to include Crustaceans, Myriapods and Spiders. The name
Crustacea and Myriapoda were first introduced by Cuvier (1769-1832) and Latreille (1825).
ADVERTISEMENTS:
Lamarck (1744-1829) in his classification, included spiders, mites, myriapods and silver fishes
under Arachnida and grouped prawns, lobsters, crabs and water fleas within Crustacea. It was
Cuvier who first suggested to include these animals and annelids under one large group,
Articulata. Von Siebold (1845) later separated the annelids and the rest were included under
Arthropoda.
Definition of Arthropods:
Bilaterally symmetrical and metamerically segmented animals; body covered with jointed
chitinous exoskeleton, moulted periodically and with jointed segmental appendages; coelom
highly reduced and haemocoelomic body cavity.
Habit and Habitat of Arthropods:
The arthropods are seen from 30,000 feet below to 20,000 feet above the sea level. These
bilaterally symmetrical, jointed-leg invertebrates may be marine, fresh-water, terrestrial,
subterranean and aerial. Some arthropods like barnacles are sedentary.
Innumerable crustaceans which live as planktons move passively in the current of water. But
well-developed structures are present in many arthropods for moving effectively by swimming,
crawling and flying.
ADVERTISEMENTS:
Some arthropods live within burrows, some are efficient diggers and many others build well-
designed nests. Certain arthropods like honey-bees, ants and termites are polymorphic and lead a
complicated social life. All the food habits—herbivorous, carnivorous and omnivorous are seen
among arthropods and various food-getting devices are met within this group.
Large numbers of arthropods live as parasites, and structural changes occur in them to adjust
with the peculiar mode of life. Many arthropods are well- known for their habit of migration.
Some of them can produce sound and nearly all are equipped with efficient sense organs.
Some forms exhibit a phenomenon—suspended animation, to overcome unfavourable con-
ditions. Sexual reproduction is often accompanied by courtship dances. The members may either
be oviparous or viviparous or ovoviviparous and some forms exhibit parental care.
Parthenogenesis is quite common in arthropods.
Characteristic Features of of Arthropods:
1. Body is bilaterally symmetrical and is metamerically segmented, coelomates.
2. Anterior segments are specialized to form a distinct head and tagmatization (body region) is
highly developed (e.g., head, thorax and abdomen).
3. Body is covered by jointed hard chitinous exoskeleton (usually composed of carbohydrate and
protein) with sclerotised plates moulted periodically.
4. Presence of paired jointed segmental appendages.
5. Presence of musculature with distinct striped muscles.
6. Body cavity or true coelom is much reduced and acts as haemocoel (blood cavity).
ADVERTISEMENTS:
7. Circulatory system is open type (e.g., blood vessels open within haemocoel) with a dorsally
placed tubular heart containing paired lateral ostia.
8. Haemocyanin is the usual respiratory pigment.
9. Nephridia are represented by the paired saccular excretory organs (e.g., coxal glands, antennal
or maxillary glands) of many arthropods. The Malpighian tubules act as excretory organs, found
in many terrestrial arthropods.
10. Central nervous system includes a dorsally placed anterior brain and ganglionated double
ventral nerve cord.
11. Presence of compound eyes in many groups (e.g., many crustaceans and most insects), in
which each eye is composed of several visual units (e.g., ommatidia).
12. Dorsal coelomic gonads.
13. Sexes are usually separate (= gonochoristic); some hermaphrodite.
14. Parthenogenesis is seen in some members of Insecta, Branchiopoda and Copepoda.
15. Fertilization internal in terrestrial species but external in aquatic species.
16. Eggs are centrolecithal.
17. Cleavage commonly superficial; a few spirals (e.g., Barnacles, copepods, cladocerans).
18. Development may be direct or indirect (e.g., via larval stages).
19. Absence of ciliated larvae.
Classification of Arthropods:
Classification in Outline:
In the studies of the middle part of 20th century, considerable rearrangement has been made in
the scheme of arthropod classification.
The classification given below is based primarily on the studies of Vandel, A (In Traite de
Zoologie, Tome VI, ed, P. Grassae pp. 79-158, 1949) and Snodgrass, R. E. (Arthropoda, in
‘McGraw-Hill Encyclopedia of Science and Technology’ Vol. I, McGraw- Hill, New York,
1960).
Phylum Arthropoda:
I. Subphylum Trilobitomorpha or Trilobita:
(i) Class Trilobita, e.g., Agnostus, Trinucleus.
II. Subphylum Chelicerata or Arachnomorpha:
(i) Class Merostomata
1. Subclass Xiphosurida e.g., Limulus, Tachypleus.
2. Subclass Eurypterida e.g., Eurypterus.
(ii) Class Arachnida
Order Scorpionida e.g., Palamnaeus, Buthus.
Order Uropygi e.g., Mastigoproctus, Trithyreus.
Order Amblypygi e.g., Sarax, Myodalis, Tarantula.
Order Palpigradi e.g., Eukoenenia, Prokoenenia.
Order Araneida e.g., Araneus, Argiope, Lycosa, Latrodectus.
Order Ricinulei e.g., Cryptocellus, Ricinoides.
Order Pseudoscorpionida e.g., Garypus, Faella.
Order Solifugae e.g., Galeodes, Eremobates.
Order Opiliones e.g., Trogulus, Mitobates.
Order Acardia e.g., Trombicula, Argas, Boophilus, Dermacentor.
(iii) Class Pycnogonida e.g., Nymphon, Pycnogonum.
III. Subphylum Mandibulata:
(i) Class Crustacea
1. Subclass Cephalocarida
Order Cephalocarida e.g., Hutchinsoniella, Lightiella.
2. Subclass Branchiopoda
Order Anostraca e.g., Artemia, Branchipus.
Order Notostraca e.g., Triops, Lepidurus.
Order Diplostraca e.g., Leptodora, Daphnia.
3. Subclass Ostracoda
Order Mydocopa e.g., Philomedes, Cypridina.
Order Cladocopa e.g., Polycope
Order Platycopa e.g., Cytherella.
Order Podocopa e.g., Cypris, Darwinula.
4. Subclass Mystacocarida e.g., Derocheilocaris
5. Subclass Copepoda
Order Calanoida e.g., Calanus, Diaptomus
Order Harpacticoida e.g., Attheyella, Harpacticus.
Order Cyclopoida e.g., Cyclops, Ergasitus
Order Notodelphyoida e.g., Natodelphys, Doropygus.
Order Monstrilloida e.g., Monstrilla.
Order Caligoida e.g., Caligus, Eudactylina.
Order Lernaeopodoida e.g., Brachiella, Lernaea.
6. Subclass Branchiura
Order Branchiura e.g., Argulus, bolops.
7. Subclass Cirripedia
Order Thoracica e.g., Balanus, Lepas.
Order Acrothoracica e.g., Cryptophialus, Trypetesa.
Order Apoda e.g., Proteolepas
Order Rhizocephala e.g., Sacculina, Peltogaster.
Order Ascothoracica e.g., Synagoga, Dendrogaster.
8. Subclass Malacostraca.
1. Super order Phyllocarida
Order Nebaliacea e.g., Nebaliopsis, Nebalia.
2. Super order Hoplocarida
Order Stomatopoda e.g., Squilla, Coronida.
3. Super order Syncarida
Order Anaspidacea e.g., Anaspides, Paranaspides.
Order Bathynellacea e.g., Bathynella.
4. Super order Peracarida
Order Mysidacea e.g., Mysis, Neomysis.
Order Cumacea e.g., Cumopsis, Diastylis.
Order Tanaidacea e.g., Tanais, Neotanais
Order Isopoda e.g., Oniscus, Ligia.
Order Amphipoda e.g., Gammarus, Caprella.
5. Super order Eucarida
Order Euphausiacea e.g., Euphausia, Nematoscelis.
Order Decapoda e.g., Palaemon, Homarus, Palinurus, Scyllarus, Hippa, Eupagurus, Cancer.
(ii) Class Chilopoda
Order Scutigeromorpha e.g., Scutigera.
Order Lithobiomorpha e.g., Lithobius.
Order Scolopendromorpha e.g., Scolopendra
Order Geophilomorpha e.g., Geophilus.
(iii) Class Symphyla e.g., Scutigerella, Scolopendrella.
(iv) Class Pauropoda e.g., Pauropus.
(v) Class Diplopoda e.g., Scutigerella, Scolopendrella.
(a) Subclass Pselaphognatha
Order Pselaphognathae e.g., Polyxenus.
(b) Subclass Chilognatha
Order Platydesmida e.g., Platydesmus.
Order Polyzoniida e.g., Polyzonium.
Order Polydesmida e.g., Polydesmus.
Order Chordeumida e.g., Chordeuma.
Order Julida e.g., julus
Order Spirobolida e.g., Spirobolus
Order Spirostreptida e.g., Thyropygus.
(vi) Class Insecta or Hexapoda
(a) Subclass Apterygota
1. Super order Entognatha
Order Protura e.g., Eosentomon, Acerentomen.
Order Collembola e.g., Isotoma, Neanura.
Order Diplura e.g., Campodea, Heterojapyx.
2. Super order Gctognatha
Order Thysanura e.g., Lepisma, Machilis.
(b) Subclass Pterygota
1. Section Paleoptera
Order Ephemeroptera e.g., Ephemera, Hexagenia.
Order Odonata e.g., Aeschna, Libellula, Ischnura.
2. Section Polyneoptera
Order Dictyoptera e.g., Periplaneta, Mantis.
Order Isoptera e.g., Termes, Odontotermes.
Order Zoraptera e.g., Zorotypus.
Order Plecoptera e.g., Perla, Isoperla.
Order Notoptera e.g., Crylloblatta.
Order Cheleutoplera e.g., Carausius, Phyllium.
Order Orthoptera e.g., Hieroglyphus, Tryxalis, Locusta Schistocerca, Gryllotalpa.
Order Embioptera e.g., Embia.
Order Dermaptera e.g., Forficula.
3. Section Oligoneoptera
Order Coleoptera e.g., Photinus, Calandra, Adalia, Dineutus.
Order Megaloptera e.g., SialiSi Corydalis.
Order Raphidioptera e.g., Raphidia.
Order Planipennia e.g., Mantispa, Myrmeleon.
Order Mecoptera e.g., Panorpa.
Order Trichoptera e.g., Rhyacophilia.
Order Lepidoptera e.g., Parides, Papilio, Bombyx.
Order Diptera e.g., Anopheles, Musca.
Order Siphonaptera e.g., Pulex, Ctenocephalus.
Order Hymenoptera e.g., Apis, Vespa, Formica.
Order Strepsiptera e.g., Stylops.
4. Section Paraneoptera
Order Psocoptera e.g. Psocus.
Order Mallophaga e.g., Menopon.
Order Anoplura e.g., Pediculus.
Order Thysanoptera e.g., Heliothrips.
Order Homoptera e.g., Cicada, Aphis, Tachardia.
Order Heteroptera e.g., Cimex, Anasa, Triatoma.]
Indicates extinct group.
The classification, as given in the text book of Parker and Haswell (1972, 7th ed.), shows that the
phylum Arthropoda is divided into seven subphyla.
The subphyla are:
Subphylum I. Onychophora
Subphylum II. Tardigrada
Subphylum III. Pentastomida
Subphylum IV. Trilobitomorpha
Subphylum V. Chelicerata
Subphylum VI. Pycnogonida
Subphylum VII. Mandibulata
The classification of Arthropoda of Ruppert and Barnes (1994) as given partly in this textbook
(4th ed.), shows that the phylum is divided into four subphyla.
They have upgraded the class Crustacea into subphylum rank and the subclasses under Crustacea
have also been upgraded into class rank. Most recent zoologists are in favour to retain Crustacea
as a class and Remipedia, Cephalocarida, Branchiopoda, Ostracoda, etc., keep them as subclasses
rank.
These subphyla include the following classes:
Remarks:
Ruppert and Barnes (1994) have placed pentastomids as a separate class Pentastomida, related to
Branchiurans and Copepods but Anderson (1998) has placed these parasitic worms in a separate
phylum Pentastomida.
Classification with Characters:
1. Subphylum Trilobita (or Trilobitomorpha) [Gk. tri = three, lobos = lobe, morphe =
shape = three-lobed form]
Characters:
1. Extinct marine arthropods.
2. Body more or less oval and flattened from above downwards.
3. Body is divided into three regions:
(i) The anterior head or cephalon,
(ii) the middle region trunk or thorax and
(iii) a posterior pygidium.
4. Each region of the body is divided into 3 lobes by two longitudinal furrows, hence the animals
derive their name Trilobites or three-lobed form.
5. Size varies from 10 mm to 60 cm.
6. Head and pygidium were covered by an un-jointed calcareous exoskeleton, called carapace.
7. Presence of a pair of compound eyes, found laterally on the anterior part of the body.
8. A pair of many-jointed antennae represents the pre-oral appendage.
9. Post-oral appendages are uniform, biramous and unspecialized. The innermost branch of each
appendage was without long setae and was probably adapted for walking and the outermost
branch had long filaments used for swimming or filtering food materials. The two branches are
sometimes called endopodite and exopodite also.
10. Each leg has 8 segments.
11. The anal opening was on the last segment of the pygidium.
The subphylum includes 4000 species which are grouped under 5 classes and the class Trilobita
includes the largest number of species.
Examples:
Agrestus, Ampyx, Mesonocis, Holmia, Trinucleus, etc.
2. Subphylum Chelicerata or Arachnomorpha [Gk. chele = claw]
Characters:
1. Heterogenous group of arthropods, in all of which pre-oral antennules or first antennae are
absent (nonantennate).
2. Body is divided into two parts— cephalothorax (or prosoma) and abdomen (or opisthosoma)
with no distinct head.
3. Cephalothorax (or prosoma) possesses five postoral segments, each with a pair of appendages.
4. First pair of appendages on the first postoral segment is called chelicerae and are feeding
appendages. The chelicerae become pre-oral in position. First pair of appendages is not antennae
but chelicerae, used in feeding.
5. Chelicerates have no jaws (mandibles); hence may be called amandibulates.
6. Each chelicera is jointed and bears a terminal chela.
7. Abdomen (opisthosoma) consists of 12-13 segments and a telson (telson and many abdominal
segments are absent in certain forms).
8. Second abdominal segment bears genital aperture which remains covered by a modified
abdominal appendage, called operculum.
9. Compound eyes in most cases degenerated.
10. Median simple eyes present.
11. Development usually direct.
12. Primarily marine arthropods, although most living forms are terrestrial.
It includes 3 classes:
(i) Merostomata
(ii) Arachnida and
(iii) Pycnogonida.
Class 1. Merostomata [Gk. meros = the thigh; stomatos, genitive of stoma = mouth]:
Characters:
1. Marine forms with fairly developed compound eyes, present laterally.
2. Head and thorax are fused into a single unit—the prosoma or cephalothorax covered by a
single sheet of exoskeleton, the carapace.
3. First pair of appendages on the prosoma is the Chelicerae followed by 5 pairs of appendages,
the walking legs.
4. Prominent caudal spine, called telson, present at the end of the body, used as a lever in
pushing and balancing during locomotion.
5. Respiratory organs are gills (book- gills), which are borne on the plate-like appendages of the
mesosoma.
6. Adults crawl on earth with the face downwards, but young can swim actively.
The class is divided into two subclasses:
(i) Xiphosura and
(ii) Eurypterida.
Subclass 1. Xiphosura [Gk. Xiphos = sword, oura = tail], Horse-shoe crab. Many fossil
forms and 4 living species.
Features:
1. Cephalothorax (prosoma) is covered by a broad, smooth, and horse-shoe shaped carapace
which is convex above and bears 2 pairs of eyes, one compound and lateral, and the other pair
simple and median in position.
2. Caudal spine is elongated, slender and pointed.
3. Dorsal ridge is visible in the abdomen.
4. Abdomen (opisthosoma) bears 5 pairs of book-gills.
5. Excretion is performed by a four-lobed coxal gland.
6. Development with a larval stage, called trilobite larva.
7. These marine, bottom dwellers are commonly called horse-shoe crabs.
Examples:
Limulus, Tachypleus, Carcinoscorpius.
Subclass 2. Eurypterida or Gigantostraca (Water scorpions)
Features:
1. All are extinct forms.
2. Scorpion-like appearance.
3. Body is compressed dorsoventrally and protected by a chitinous exoskeleton.
4. Short prosoma is covered by dorsal carapace and consists of 6 segments fused together.
5. Trunk or abdomen is followed behind prosoma and consists of 12 free segments.
6. At the end of the trunk or abdomen there is a post and tail-plate or spine which may be
triangular (e.g., Eurypterus) or divided into two lobes (e.g., Pterygotus).
7. Mouth ventrally placed of the prosoma.
8. Five pairs of prosomal appendages except 1st pair are 6-8 joints each and are associated with
locomotion.
Examples:
Euryptarus (Ordovician- Permian), Pterygotus (Ordovician), Slimonia (Silurian).
Class 3. Arachnida [Gk. arachne = spider] Approx. 74,000 species.
Characters:
1. Body divided into two regions— cephalothorax (Prosoma) and abdomen.
2. Eyes usually simple.
3. Compound eyes when present are degenerated.
4. Two pairs of jointed cephalic append- ages-chelicerae and pedipalpi present. The first pair of
cephalic appendages, known as chelicerae, which are preoral and the 2nd pair, the pedipalps, are
postoral and serve partly as jaws.
5. Four pairs of thoracic legs present.
6. Abdominal segments often reduced and abdominal appendages not associated with
locomotion.
7. Antennae absent.
8. In terrestrial forms, the respiratory organs are book-lungs or tracheae or both in some species.
9. Excretory organs are malpighian tubules or coxal glands or both.
10. Sexes separate.
11. Eggs yolky and centrolecithal.
12. Development not accompanied by metamorphosis.
13. Carnivorous and mostly terrestrial.
The class Arachnida is divided into 10 orders:
These are:
(i) Scorpionida,
(ii) Uropygi,
(iii) Amblypygi,
(iv) Palpigradi,
(v) Araneida,
(vi) Ricinulei,
(vii) Pseudoscorpionida,
(viii) Solifugae,
(ix) Opiliones and
(x) Acarida.
Order 1. Scorpionida:
1. These are terrestrial arachnids found under legs, stones or in rock crevices of tropical and
subtropical regions.
2. Body is divided into prosoma (cephalothorax) and opisthosoma (abdomen).
3. Prosoma unsegmented and covered by carapace bearing a pair of chelicerae, a pair of
pedipalpi and four pairs of walking legs.
4. Abdomen divisible into two parts— broad pre-abdomen (mesosoma) consisting of 7 segments
and narrow post- abdomen (metasoma) consisting of 5 segment with a caudal spine formed by
the modification of telson.
5. Chelicerae are small 3-segmented but pedipalpi are large and 6-segmented.
6. Second segment of the pre-abdomen bears two comb-shaped pectines.
7. Four pairs of respiratory organs, called book lungs.
8. Viviparous.
This order includes scorpions.
Examples:
Palamnaeus, Mesobuthus, Buthus, Scorpio, etc.
Order 2. Uropygi:
1. These are commonly called whip scorpions.
2. Size ranges from 2-65 mm in length.
3. Telson is present as a long whip-like flagellum at the posterior end.
4. Poison glands are absent but an acidic substance from the anus is sprayed.
5. Female provided with a brood sac to carry eggs.
6. Upper lip of the rostrum and the bases of the pedipalps form a peculiar ball and socket joint
around the mouth to act as a filtering apparatus.
7. Short and stout pedipalp.
8. Second leg is antenna-like and elongated.
9. Abdomen bears 12 segments.
10. Book lungs placed on the second and third abdominal segments,
11. Nocturnal and carnivorous.
Examples:
Mastigoproctus, Trithyreus.
Order 3. Amblypygi:
1. Commonly known as tailless whip scorpions.
2. Size varies from 4-45 mm in length.
3. Body flattened dorsoventrally.
4. Telson almost absent.
5. Abdomen consists of 11 segments.
6. Chelicerae are 2-jointed and hook-like.
7. Pedipalps consist of seven segments and are stout and raptorial.
8. A prominent movable claw is present at the distal end of each pedipalp.
9. Book lungs open on the ventral side of second and third abdominal segments.
10. Females carry brood-sacs during breeding season.
Example:
Tarantula.
Order 4. Palpigradi:
1. Commonly called microwhip scorpions.
2. Size ranges from 0.5-3 mm in length.
3. In the cephalothorax last two segments are not united.
4. Telson with a many-jointed flagellum.
5. Chelicerae are 3-segmented and chelated.
6. Pedipalpi leg-like.
7. Both simple and compound eyes are absent.
8. Second pair of legs act as antennae.
9. Respiration is cutaneous.
10. In addition, paired eversible sacs in the abdominal segments also act as respiratory organs.
Example:
Eukoenenia.
Order 5. Araneida:
1. Commonly called spiders.
2. Cephalothorax un-segmented.
3. Abdomen in general is also un-segmented, soft and round.
4. Four pairs of eyes are present.
5. Chelicerae are complex and poison glands open through them.
6. Pedipalpi simple and six-jointed.
7. Book lungs are associated with trachea for respiration.
8. Spinning glands usually present and appendages of fourth and fifth abdominal segment form
spinnerets.
Examples:
Argiope, Aranea (orb-web spider), Latrodectus, Lycosa (wolf-spider), etc.
Order 6. Ricinulei:
1. Commonly called Ricinuleids.
2. Sizes usually less than 10 mm in length.
3. Body short and compact.
4. Cephalothorax is drawn anteriorly into a many-jointed movable projection, called cucullus.
5. Eyes are absent.
6. Abdomen superficially looks like four segmented but truly it consists of nine segments.
7. Both the chelicerae and pedipalpi are chelated.
8. Tracheae are the respiratory organs.
Examples:
Cryptocellus, Ricinoides.
Order 7. Pseudoscorpionida:
1. These are commonly called pseudoscorpions.
2. Size is never more than 8 mm in length.
3. Body oval and flattened dorsoventrally.
4. Abdomen broad and consists of eleven segments.
5. Abdomen is not separated into pre- and post-abdomen and does not bear the caudal sting.
6. Simple eyes are absent.
7. Chelicerae are small and contain silk glands.
8. Pedipalpi is scorpion-like and contains poison glands.
9. Respiratory organs are tracheae.
10. Malpighian tubules are absent.
11. Oviparous.
12. These are found under the bark of trees.
Examples:
Chelifer, Neobisium.
Order 8. Solifugae:
1. Commonly known as wind scorpions.
2. Size varies form 1-7 cm in length.
3. These are swift runners.
4. Body is divided in 3-regions—head, thorax and abdomen.
5. Abdomen oval and contains ten segments.
6. Chelicerae large and prominent.
7. Pedipalpi elongated and resembles the legs.
8. Poison glands are absent.
9. Respiratory organs are well-developed tracheae.
10. All are nocturnal, ferocious and predators.
Examples:
Galeodes, Eremobates.
Order 9. Opiliones:
1. Commonly called harvestmen.
2. Size varies from 1-22 mm in length.
3. Cephalothorax un-segmented.
4. Abdomen consists of 10 segments and is not distinctly separated from the cephalothorax.
5. Telson absent.
6. Compound eyes are lacking.
7. Pedipalpi is leg-like.
8. Walking legs are usually very long and slender.
9. Respiration takes place through tracheae.
10. No poison or spinning glands.
11. Males possess copulatory organ and females are provided with ovipositor.
Examples:
Caddo, Phalangium, Trogulus, Mitobates.
Order 10. Acarida:
1. Commonly known as ticks and mites.
2. Number of forms is microscopic in sizes.
3. Body without any external division.
4. Chelicerae and pedipalpi are usually small and associated with the mouth- parts which are
adapted for—biting, piercing or sucking.
5. Chelicerae are usually composed of 2 to 3 segments but may have up to 6 segments.
6. Chelicerae may be pincer-like, fanglike or lance-like.
7. Pedipalpi also may be variously modified.
8. Legs are provided with claws.
9. Respiratory organs are tracheae or in many, respiration is cutaneous.
10. Most of them are parasites on man and other animals.
11. Some are notorious pests of agricultural products.
Examples:
Tetranychus, Demodex, Hoplophorella, Annectacarus, Sarcoptes.
Class 3. Pycnogonida (= Pantopoda):
Approx. 1,000 species; about 16 Indian species.
Characters:
1. Partially sedentaric marine chelicerates, commonly called sea spiders.
2. Young’s are parasitic on different soft bodied invertebrates.
3. Externally segmented body.
4. Reproductive openings present on the leg segments and not abdominal.
5. Chelicerae short and pedipalpi segmented.
6. Head with proboscis.
7. Third pair of appendages in the male carries the eggs and is called the ovigers.
8. Trunk of 3-6 segments with long walking legs.
9. Opisthosoma much reduced with a terminal anus.
10. Protonymphon larva in the life cycle of most pycnogonids.
Examples:
Nymphon, Pycnogonum, Colossendeis.
3. Subphylum Crustacea:
[L. crusta – a hard shell] Copepods, shrimps, prawns, barnacles, lobsters crabs. Approx.
45000 species; about 3000 Indian species.
Characters:
1. Body divisible into 3 regions—head, thorax and abdomen.
2. Two pairs of antennae are a distinguishing feature among crustaceans.
3. Other cephalic appendages are a pair of mandibles and two pairs of maxillae.
4. Thoracic and abdominal appendages are usually 8 pairs and 6 pairs, respectively, variable in
lower crustacea.
5. Appendages typically biramous except of antennules.
6. Carapace covers all or part of the body.
7. Head bears a pair of compound eyes on movable jointed stalk.
8. Respiration takes place either by gills or by the general surface of the body when the
exoskeleton is thin or by some of the limbs.
9. Vascular system consists of a contractile heart, arteries and haemocoelomic spaces.
10. Excretory organs are the modification of coelomoducts may be either antennal glands (green
glands) or shell glands (maxillary glands found in the second pair of maxillae).
11. Brain formed by the fusion of first four embryonic ganglia and is connected with ventral
nerve cord by oesophageal connectives.
12. Sexes separate.
13. Distinct sexual dimorphism present.
14. Eggs usually centrolecithal, i.e., yolk present in the central part of the egg, or may be
telolecithal, i.e., yolk occupies one-half of the egg, or alecithal, i.e., without yolk.
15. Development includes a larval form, the nauplius, bearing a single median eye and 3 pairs of
appendages.
16. Mainly aquatic, mostly marine, many freshwater and some have invaded into terrestrial
condition.
The subphylum crustacea is divided into 11 classes:
(i) Remipedia,
(ii) Cephalocarida,
(iii) Branchiopoda,
(iv) Ostracoda,
(v) Copepoda
(vi) Mystacocarida,
(vii) Branchiura,
(viii) Pentastomida,
(ix) Tantulocaride,
(x) Cirripedia and
(xi) Malacostraca.
Remarks:
Many zoologists, such as Ruppert and Barnes (1994) use the category subphyla:
(i) Crustacea for typical bi-ramous appendages (e.g., copepods, barnacles, shrimps, lobsters and
crabs) and
(ii) Uniramia for uniramous appendages (e.g., centipedes, millipedes and insects) instead of
subphylum Mandibulata.
Pechenik (2000) and many zoologists agreed to use Mandibulata containing the classes—
Crustacea, Myriapoda and Insecta.
He used Myriapoda (Gk. many feet) containing 4 orders, such as:
(i) Order Chilopoda (e.g., centipedes),
(ii) Order Diplopoda (e.g., millipedes),
(iii) Order Symphyla (e.g., symphylans) and
(iv) Order Pauropoda (e.g., pauropods).
The class Myriapoda is characterised by many segmented trunk, with each bearing uniramous
legs, a pair of antennae, compound eyes absent and malpighian tubules for excretion.
Class 1. Remipedia:
This group was first recognised in 1983 with twelve known species.
Features:
1. Small and worm-like bodies, range up to 30 mm in length.
2. Head covered by a head-shield, followed by a trunk of 20-30 similar segments.
3. Each segment of the body bears a pair of lateral biramous appendages.
4. Telson with caudal rami.
5. Hermaphrodite.
They are the inhabitants of tropical marine caves.
Examples:
Lasionectes, Speleonectes.
Remarks:
Morphological data of this group suggest to be the closest of all the animals to the ancestral
crustacean body but molecular data remain ambiguous.
Class 2. Cephalocarida [Cephalocarids; Approx. 9 species]:
The members of this group are considered to be most primitive among living crustaceans and the
first member was discovered in Long Island Sound in 1955. The all species are marine and have
collected in the soft sediments of the bottom up to the depths of over 1,500 m.
Features:
1. Small-sized animals exceeding 3.7 mm in length.
2. Horse-shoe shaped head followed by an elongated trunk.
3. First 8 trunk segments bear biramous appendages which are identical in appearance.
4. The appendages are tripartite.
5. Exopodites of these appendages are four-jointed and leaf-like and bear lateral
pseudoepipodite.
6. Endopodites are segmented, cylindrical and ambulatory in function.
7. Movements of the limbs produce water current for locomotion and also for collecting food.
8. Eyes are buried in the head.
9. Hermaphrodite.
Examples:
Hutchinsoniella, Lightiella.
Class 3. Branchiopoda (Gk. branchiona fin); Approx. 800 species:
Features:
1. Mostly freshwater species, a few are marine.
2. Trunk appendages are uniform and leaf-like.
3. Presence of one pair un-jointed or jointed caudal styles.
4. Carapace either absent or shield-like or bivalve.
5. First antennae and maxillae are small and in some cases absent.
6. Mandibular palp either rudimentary or absent.
It has 3 living orders:
(i) Anostraca
(ii) Notostraca and
(iii) Diplostracs.
Order (i) Anostraca:
1. It includes forms like fairy shrimps and brine shrimps.
2. Carapace is absent.
3. They possess stalked eyes.
4. Antennae redudced and triangular in females.
5. In males, the antennae are stout copulatory structures.
6. Trunk is elongated and first 11 segments bear alike legs.
7. Caudal styles un-jointed.
Examples:
Branchipus, Artemia.
Order (ii) Notostraca:
1. Commonly called Tadpole shrimp.
2. Carapace is large and shield-shaped.
3. Possesses 35-71 pairs of legs.
4. Eyes are sessile and placed close together.
5. Antennae rudimentary.
6. Caudal styles filamentous and many- jointed.
7. Parthenogenesis is frequent.
Examples:
Triops, Lepidurus.
Order (iii) Diplostraca:
1. It includes forms like clam-shrimps (Estheria) and water fleas (Daphnia, Leptodora).
2. Carapace laterally compressed, bivalved and enclosing trunk.
3. Biramous and large antennae are used during swimming.
4. Caudal styles clawed and un-jointed.
Examples:
Daphnia, Sida, Leptodora.
Class 4. Ostracoda (Gk. ostrakodes – testaceous resembling a shell); Approx. 7000 species:
Features:
1. Small crustaceans and commonly called seed-shrimps.
2. Mostly marine or freshwater, a few in terrestrial habitats.
3. Body enclosed in a hinged bivalved carapace.
4. Trunk appendages never more than 2 pairs.
5. Mandible with a palp.
6. Both pairs of antennae modified for swimming.
7. Respiration usually cutaneous.
8. Eyes may or may not be present.
9. Males are rare and the second antennae of the males serve as clasping organs.
The class Ostracoda includes 4 orders:
(i) Mydocopa
(ii) Cladocopa
(iii) Platycopa and
(iv) Podocopa.
Order (i) Mydocopa:
1. Carapace bears an aperture through which the antennae protrude.
2. Antenna is operated by powerful muscles.
3. Limbs are four pairs.
4. Heart with paired ostia.
5. Sessile compound eyes.
Examples:
Philomedes, Cypridina.
Order (ii) Cladocopa:
1. The aperture for the protrusion of antennae is absent.
2. Each second antenna bears two branches or rami.
Example:
Polycope.
Order (iii) Platycopa:
1. Antennal aperture absent.
2. Rami of second antennae broad.
3. Limbs three pairs.
Example:
Cytherella.
Order (iv) Podocopa:
1. Antennal aperture on the carapace absent.
2. Endopodite of the second antenna well developed than exopodite and carries a claw.
3. Mandibular palp composed of four articles.
4. Four pairs of limbs are posteriorly placed.
Examples:
Cypris, Darwinula.
Class 5. Copepoda (Gk. kope = handle) Approx. 8,500 known species:
Features:
1. Mostly small crustaceans.
2. Body with well-marked segments.
3. Trunk composed of a thorax bearing 5 pairs of biramous appendages used for swimming.
4. Abdomen without appendages.
5. Presence of a pair of caudal styles.
6. Head-shield present but no carapace.
7. Single median nauplius eye present but paired compound eyes absent.
8. Well-developed antennae may or may not be used for swimming.
9. Seventh segment of the body bears the reproductive apertures.
It includes 7 orders:
(i) Calanoida
(ii) Harpacticoida
(iii) Cyclopoida
(iv) Notodelphyoida
(v) Monstrilloida
(vi) Caligoida and
(vii) Lernaeopodoida.
Order (i) Calanoida:
1. Free-living and size is fairly large.
2. Posterior part of the trunk is distinctly separated from the anterior part and between genital
and pregenital segments.
3. First antennae of female have 23-25 segments.
4. The second antenna is biramous.
5. The female possesses a single median egg-sac.
Examples:
Canthocalanus, Eucalanus, Calanus, Paracalanus, Acrocalanus, Centropages, Phyllodioptomus,
Heliodioptomus, Allodioptomus, Temora, Candacia, Calanopia, Acardia, Tortanus, Diaptomus.
Order (ii) Harpacticoida:
1. Free-living.
2. Trunk not constricted in the middle.
3. First antenna in the female is short and consists of 5-9 segments.
4. Second antenna is biramous.
Example:
Attheyella.
Order (iii) Cyclopoida:
1. Free-living.
2. Posterior part of the trunk is separated from the anterior part and it includes pregenital
segments.
3. First antenna of female has more than 17 articles.
4. Second antenna is uniramous.
5. Female carries a pair of egg-sacs.
6. Some members are parasitic.
Examples:
Cyclops, Ergasilus, Eucyclops.
Order (iv) Notodelphyoida:
1. Only in males the posterior half of the trunk bears pregenital segments.
2. Both sexes live as commensal within a tunicate, Ascidia.
Example:
Notodelphys.
Order (v) Monstrilloida:
1. Free-swimming and completely marine.
2. Adults without mouth parts, antennae and alimentary canal.
3. Larva starts as a free-swimming nauplius and parasitises a mollusc or annelid.
4. It draws nourishment by its antennae from the host.
5. Adult, when full grown, becomes again free-swimming and perform only reproduction.
Example:
Monstrilla.
Order (vi) Caligoida:
1. Lives as ectoparasite on fishes.
2. Posterior part of trunk carries two pregenital segments.
3. The antennae are modified to act as adhesive organs.
Examples:
Caligus, Eudactylina.
Order (vii) Lernaeopodoida:
1. Adults remain as ectoparasites up to the attainment of sexual maturity.
2. The sexually matured forms become free-swimming.
3. After copulation, female again becomes parasitic on fishes.
4. It changes into a worm-like form and carries the eggs in a brood chamber.
5. Larvae start as free-swimming nauplius but soon infect a new host.
Examples:
Lernaea, Lernaeocera.
Class 6. Mystacocarida —Approx. 8 known species:
This subclass was created after the discovery of several crustaceans in the year in 1943.
Features:
1. Marine and interstitial.
2. Length of the body always within 1 mm.
3. Cylindrical bodies with distinct cephalic appendages.
4. Trunk consists of 5 segments each with a pair of appendages.
5. Nauplius eye persists and the compound eyes absent.
6. Two caudal styles work as pincers.
Examples:
Derocheilocaris.
Class 7. Branchiura —Approx. 130 known species:
Features:
1. Fish ectoparasites.
2. Dorsoventrally flattened body with suctorial mouth.
3. Broad shield-like carapace covers the cephalothorax.
4. Small, un-segmented and bilobed abdomen.
5. Sessile compound eyes present.
6. Flagella present in the appendages of some body segments.
7. 5 pairs thoracic appendages.
8. Fifth body segment bears the genital apertures.
9. Males have two testes but females possess a single ovary.
10. Commonly called fish lice.
It includes a single order, having the same name and includes a single family.
Examples:
Argulus, Dolops.
Class 8. Pentastomida—Approx. 100 known species:
Features:
1. All the members are parasitic and live mainly in the lungs and nasal passages of reptiles, but
some species parasitize amphibians, birds and mammals including dogs and man.
2. Worm-like body ranges 2 to 13 cm long, of which the females are 10 cm in length.
3. Larva possesses 2-3 pairs of un-jointed Legs.
4. Adults are legless but possess only 4 pairs of anterior chitinous hooks, used for clinging to the
host tissues.
5. Body covered by a non-chitinous cuticle and exhibits annular markings over the abdomen in
the adult.
6. Exoskeleton moulted periodically.
7. Muscles are striated and metamerically arranged.
8. Most of the systems, such as digestive, excretory and reproductive are modified to adapt the
endoparasitic life.
9. Completion of the life history requires intermediate host.
10. They are gonochoristic, i.e., the sexes are separate.
11. Fertilization internal.
12. Cleavage spiral.
13. Pentastomids are popularly known as Tongue worms or sometimes referred to as “five
mouths”.
Remarks:
The taxonomic status of Pentastomids has long been uncertain. Previously this group was treated
as a separate phylum. But recently the sperm ultrastructure and analyses of DNA sequences
coding for 18s ribosomal RNA indicate the similarities with crustaceans and suggest that
Pentastomids are closely related to marine crustaceans, specially with branchiurans and
copepods.
Examples:
Raillietiela, Cephalobaena, Linguatula, Armillifer.
Class 9. Tantulocarida—Approx. 5 known species:
Features:
1. Ectoparasite on other deep water crustaceans.
2. Deep water crustaceans.
3. Appendages absent in trunk segments.
Examples:
Basipoplella.
Class 10. Cirripedia (L. cirrus = curled, pedis = foot). Approx. 1000 known species:
Features:
1. All are marine.
2. Adults are sedenteric.
3. Body poorly segmented.
4. Six pairs biramous filamentous appendages present.
5. Abdomen almost absent, with only a pair of caudal style.
6. Body enclosed within a bivalve carapace with calcareous plates on it.
7. Adults without eyes and antennae.
8. Usually hermaphrodite.
9. Young passes through nauplius and cypris stage.
10. Commonly called barnacles.
This subclass includes 5 orders:
(i) Thoracica
(ii) Acrothoracica
(iii) Apoda
(iv) Rhizocephala and
(v) Ascothoracica
Order (i) Thoracica:
1. Adults are permanently fixed by its preoral region on substrates at inter- tidal level.
2. Presence of six pairs of appendages in the trunk called cirri.
3. Abdomen without segments.
4. Some forms have a stalk, others are sessile.
Examples:
Examples are Lepas (Goose barnacles), Balanus (Acorn barnacles).
Order (ii) Acrothoracica:
1. Sessile forms and are fixed on the shell of molluscs.
2. Mantle devoid of calcareous plates.
3. Presence of only four pairs of appendages in the trunk.
4. Male lives as parasite within female.
Example:
Alcippe.
Order (iii) Apoda:
1. Completely parasitic forms.
2. Small distinctly segmented body.
3. Trunk appendages absent.
Example:
Proteolepas.
Order (iv) Rhizocephala:
1. Completely parasitic.
2. Mantle remains, but the calcareous shells are absent.
3. Body absolutely degenerated in adults due to the loss of alimentary canal and appendages.
4. No trace of segmentation.
Examples:
Sacculina, Peltogaster.
Order (v) Ascothoracica:
1. These are endoparasites on Anthozoa and Echinodermata.
2. Mantle bilobed or sac-like but plates are absent.
3. Oral appendages are modified for piercing and sucking.
4. Presence of 6 pairs of appendages in the trunk.
Examples:
Symagoga, Laura, Dendrogaster.
Remarks:
Recent trend of the crustacean classification shows that the subclasses Mystacocarida, Copepoda,
Branchiura, Tantulocarida and Cirripedia are included under the class Maxillopoda for the
characteristic features—6 thoracic and 5 abdominal segments and the first pair of trunk
appendages are maxillipeds.
Class 11. Malacostraca (Gk. malakos = soft + ostracon = a shell). Over 20,000 species:
Features:
1. Body consists of 20-21 segments.
2. Thoracic and abdominal appendages distinct from one another.
3. Carapace covers the head and at least some thoracic segments.
4. Mandible with a palp.
5. Presence of compound eyes on stalk.
6. Antennule with two-many-jointed flagella.
7. Male and female gonopores on the bases of 8th and 6th thoracic appendages.
The class includes five super orders:
(i) Phyllocarida
(ii) Hoplocarida
(iii) Syncarida
(iv) Peracarida and
(v) Eucarida.
Super order (i). Phyllocarida (Gk. phyllon = a leaf, L. caridis, genitive of caris = a shrimp):
Features:
1. A movable rostrum present at the anterior end of cephalothorax.
2. Carapace large, bivalved and encloses both the cephalothoracic and abdominal segments.
3. Thoracic legs are all alike and folia- ceous.
4. Abdomen has 7 segments.
5. Well-developed appendages in the first four abdominal segments.
6. Telson with a movable caudal furca.
It includes only one order Nebaliacea.
Order Nebaliacea:
Presence of seven abdominal segments. Telson with a pair of caudal styles. A prominent
carapace covers almost the entire length of the body. Thoracic appendages are leaf-like and serve
as respiratory surface. Abdominal appendages are biramous. The common example is Nebalia.
Super order (ii). Hoplocarida (Gk. hoplon = a weapon):
Features:
1. All are marine.
2. Carapace flat, shield-shaped and encloses the second thoracic segment.
3. Two movable segments lie anterior to the carapace.
4. Anterior one bears stalked eye and the posterior one carries the antennules.
5. Each antennule has three rami.
6. Antenna smaller than antennule.
7. Second pair of thoracic appendages is raptorial and bear a blade-like edge at its distal end.
8. Biramous abdominal appendages.
It includes only a single order Stomatopoda.
Order Stomatopoda:
Last four thoracic segments are free from the carapace. Two movable segments are present in
front of the head. Gills are carried by the abdominal appendages. Gastric glands extend up to the
telson. Heart extends up to abdomen and has thirteen pairs of ostia. The well-known examples
are Scjuilla, Pseudoscjuilla and Coronida.
Super order (iii). Syncarida (Gk. syn = together):
Features:
1. Carapace absent.
2. Elongated and tube-like heart.
3. Only the first pair of thoracic appendages modified as maxillipeds, rest are alike.
4. Biramous thoracic appendages.
5. Gills present on the thoracic appendages excepting the last one.
6. Last pair of abdominal appendages, called uropods, is fan shaped.
It includes two orders:
(i) Anaspidacea and
(ii) Bathynellacea.
Order (i) Anaspidacea:
1. First thoradc segment is fused with head.
2. They are inhabitants in the freshwater of ponds, streams of Australia, New Zealand and South
America.
Examples:
Anaspides, Allanaspides.
Order (ii) Bathynellacea:
1. They are the smaller than the members of Anaspidacea.
2. First thoracic segment is not fused with the head.
3. They are living in freshwater sediments and are world-wide in distribution.
Example:
Bathynella.
Super order (iv). Peracarida (Gk. pira = a pouch):
Features:
1. Carapace may or may not be present. When present, carapace never covers last four thoracic
segments.
2. Coxopodites of thoracic appendages bear a brood-pouch in females.
3. Presence of a tube-like, elongated heart.
4. Development direct.
This super order includes 5 orders:
(i) Mysidacea
(ii) Cumacea
(iii) Tanaidacea
(iv) Isopoda and
(v) Amphipoda.
Order (i) Mysidacea:
1. First antennae are biramous.
2. Second antenna is with scale-like squama.
3. Possesses filter feeding mechanism.
4. First pair of thoracic appendages is modified as maxillipeds.
5. Carapace is the chief respiratory surface.
6. Broad tail fin is formed by flat uropods and telson.
7. Heart is elongated but extends up to thorax and has two pairs of ostia. The common Examples
are Mysis, Hemimysis.
Order (ii) Cumacea:
1. Cephalothorax is posteriorly narrow.
2. Carapace is drawn out anteriorly to form rostrum and ventrally to form gill-chamber.
3. Sessile eyes are usually fused to form a single eye.
4. Antennae are usually un-segmented.
5. Second antennae are without exopodites and well developed in males than in females.
6. Abdomen is slender and segmented.
7. Abdominal appendages are absent in the female.
8. Uropods are rod-shaped and thus fan- shaped tail fin is absent.
The well-known examples are Cumopsis, Diastylis and Pseudocuma.
Order (iii) Tanaidacea:
1. Carapace covers first two thoracic segments.
2. Eyes, when present, are mounted on immovable stalks.
3. Second thoracic appendages are large and chelated.
4. A small squama may be present with the second antenna.
5. Uropods are slender.
The examples are Tanais, Apseudes and Neotanais.
Order (iv) Isopoda:
1. It includes aquatic, terrestrial and parasitic forms.
2. Carapace is absent, only first thoracic segment is fused into head.
3. Eyes are either without stalk or they are carried on small immovable processes.
4. Body is dorsoventrally flattened.
5. Antennule is small and rudimentary.
6. First pair of thoracic appendages is modified as maxillipeds, while the others are alike.
The well-known examples are Anthura, wood borer (Limnoria), the ectoparasite of the fish, the
Cymothoidae; parasites on the gills of shrimps and crabs (Bopyrus), wood lice (Ligia, Liriopsis,
Oniscus and Tylos, Adinda, Agnara, etc.).
Oniscus is an example of terrestrial crustacea). The wood boring isopods, Limnoria lignorum,
make deep funnels in wooden harbour for sheltering their teeming animals. They can also
damage the submarine cables.
Order (v) Amphipoda:
1. Body is flattened laterally.
2. Carapace is absent.
3. Antennules are well developed and biramous.
4. Second and third pairs of thoracic appendages are prehensile structures, called gnathopods.
5. Some thoracic appendages bear gills at their bases.
6. Abdominal appendages are of two distinct morphological forms.
The examples are Gammarus, Caprella.
Super order (v). Eucarida (Gk. eu = true, L. caridis, genitive of caris = a shrimp):
Features:
1. Carapace covers head and all the thoracic segments.
2. Mandible without sharp blade.
3. Stalked eyes.
4. Small bag-like heart, placed on the dorsal side of the thorax.
Eucarida has two orders:
(i) Euphausiacea and
(ii) Decapoda.
Order (i) Euphausiacea:
1. Thoracic appendages do not form maxillipeds and are all alike.
2. Single gill is present at the base of each thoracic appendage.
3. Pleopods are flattened.
4. In the males first two pairs are modified for copulation.
5. Elongated telson bears a movable large spine. Uropods are elongated.
The examples are krill (Euphausia, Nematoscelis, etc.), and all the members are marine, pelagic,
shrimp-like animals, and have a world-wide distribution. In the Southern Ocean they constitute a
major food source of Baleen whales.
Order (ii) Decapoda (Gk. deka = ten, podos = foot):
1. Three maxillipeds are formed by the modification of first three thoracic appendages.
2. Three sets of gills are present which differ in their arrangements.
This order is divided into 2 suborders:
(i) Dendrobranchiata and
(ii) Pleocyemata.
Suborder Dendrobranchiata:
Gills are dendrobranchiate type. The body is almost laterally compressed. The examples are
penaeid shrimps (e.g., Penaeus, Funchalia, Parapenaeus, Metapenaeopsis, Metapenaeus),
sergestid shrimp (Sergestes, Lucifer, Acetes).
The penaeid shrimps (fam. Penaeidae) are characterised by the well-developed and toothed
rosturm, carapace without postorbital spine, 3rd and 4th pairs of pleopods biramous and telson
sharply pointed with or without spines. They are found in sandy, mud estuaries, back water and
near shore areas. The penaeids are the most valuable commercial shrimps exploited in many
parts of the world.
The monophyletic theory is based on the segmentation between the annelids and arthropods, and
the prostomium and pygidium of the annelids correspond to the acron and telson of arthropods.
A number of anatomical features, such as chitinous cuticle, haemocoel, dorsal blood vessel,
segmental jointed appendages and centrolecithal eggs constitute the basis for the view that the
arthropods are monophyletic (Anderson, 1998).
The polyphyletic theory is proposed by Tiegs and Manton (1958).
They suggested that there were two ancestors of arthropods:
(i) Lobopod-annelids and
(ii) Proto- annelids (Fig. 18.121).
(i) The lobopod-annelid is considered to be the ancestor of Onychophora which was originated in
the Pre-Cambrian period about 525 million years ago. The group Uniramia which includes
chilopodes, diplopodes and insects is thought that they have evolved from the base of
onychophores.
(ii) The protoannelid ancestor is considered as the primitive annelids from which the most
primitive extinct arthropod group- Trilobitomorpha arose, from which two extant groups—
Crustacea and Chelicerata evolved by two separate evolutionary lines. Manton (1973), Ruppert
and Barnes (1994), support the polyphyletic theory.
They suggest that 3 different living groups-Chelicerata (e.g., horse-shoe crabs, spiders,
scorpions, etc.), Crustacea (e.g., copepods, barnacles, prawns, crabs, etc.) and Uniramia have
evolved separately, possibly from the annelid ancestors. This view is based on the locomotory
structures and embryonic development of the different major groups.