Phylum Arthropoda

Introduction Arthropods are said to be the most successful of all animal groups because of their huge population and their diversity. They are great in number since they can readily adapt to their particular kind of environment or habitat - be it aerial, terrestrial, or aquatic. There are two theories regarding the development and evolution of arthropods. One theory proposes that arthropods have risen from Polychaete stock while another theory states that arthropods and annelids came from a common stock. Brief History Fossil records of almost all animal phyla including Arthropoda, Cnidaria, Annelida, Echinodermata and Mollusca first appeared 550 million years ago, at the beginning of the Paleozoic Era. This is the starting point for most of the animal fossil record. Yunnan, a province in China is one example where fossil deposits have been found. Linnaeus first designated all arthropods as belonging to the term Insecta. But differences and variation in morphological and anatomical features divided this big group into different major groups of arthropods. Hence, Myriapoda which includes the centipede and millipede, Crustacea and Arachnoidea emerged. In 1845, Phylum Arthropoda included all of these mentioned groupings. Definition The arthropods are animals with segmented/ jointer appendages covered with a chitinous exoskeleton. They are schizocoelomate, triploblastic, and bilaterally symmetrical. They are considered the most successful animal group for the reason that they are great in number and they can thrive in air, water and land, with the latter being the most suitable environment. General Morphology and Physiology I. External Features a. Metamerism - evident in the embryonic development of all arthropods and is conspicuous feature of many adults especially the higher primitive species b. Tagmatization - resulted from the fusion of metamere, with the last 5 segments fused into a head, the nest 3 into the thorax with most of the abdominal segments retained. variations in tagmatization size: ! cephalothorax- fusion of head and thorax ! opisthosoma- fusion of thorax and abdomen c. eight segments of jointed appendages of arthropods c.1. coxa- most proximal c.2. trochanter c.3. femur-usually longest c.4. patella c.5. tibia c.6. metatarsus c.7. tarsus c.8. pretarsus- most distal and usually modified for a specific function II. Exoskeleton - chitinous exoskeleton secreted by the hypodermis - composition; thin outer epicuticle proteins & wax thicker inner procuticle made up of: ! outer exocuticle - made up of chitin & protein forming a complex glycoprotein, can be impregnated with salts & tannins (tanned) ! inner endocuticle- same as above but not tanned

- makes movement possible by the division of the cuticle into separate plates - articular membrane - primitively connects the plate of one segment to the plate of the adjoining segment; a region in which the cuticle is very thin and flexible - four primary plates of the cuticle of each segment: i. a dorsal tergum ii. two lateral pleura iii. a ventral sternum - joints tube-like segments or sections of the cuticular skeleton of the appendages, connected by an articular membrane; enable the segments of the appendages and body to move - pores generally penetrates the body, where secretions pass through - structures originating from the ectoderm and lined by the exoskeleton: i. tracheal tube (insects, chilopods, diplopods) ii. book lungs (scorpions and spiders) iii. proctodeum and stomodeum iv. certain portions of the reproductive tract - color depends on deposition of brown, yellow, orange or red melanin pigments within the cuticle; also present are iridescent greens, purples and other hues due to striations of the epicuticle - subcuticular chromatophores, blood and tissue pigments are other sources of pigments - ecdysis - molting which allows the arthropod to grow in size i. hypodermis detaches from the cuticle and secretes a new epicuticle ii. hypodermis secretes chitinase and proteases that seep through the newly formed epicuticle and erode away the untanned endocuticle of the old skeleton iii. hypodermis secretes a new procuticle iv. old exoskeleton splits along certain predetermined lines usually along the mid-dorsal and mid-ventral lines, and the arthropod pulls out of the encasement - after molting, the new cuticle is soft and pliable, and is stretched to accommodate the increase in size of the arthropod due to internal pressure, which in turn is due to tissue growth, and sudden uptake of air and/or water - additional procuticle may be added - instars - stages in between molts, the number of which may be fixed or continuous throughout life - ecdysomes (hormone which controls molting) III. Endoskeleton - in folding of the procuticle that produces inner projections or apodemes on which the muscles are inserted - may involve the sclerotization of internal tissue forming free plates for muscle attachment within the body IV. Muscular System - striated muscle bundles attach to the inner surface of the procuticle by tonofibrils - innervated by neurons which may be: i. phasic (fast) ii. tonic (slow) iii. inhibitory (prevents contraction) V. Coelom - schizocoelom hemocoel that are sinuses or spaces in the tissue filled with blood, lined with cilia - reduced to a cavity of the gonads or by the excretory organs VI. Circulatory System - open blood vascular system i. heart consists of one or more chambers with vascular walls arranged in a linear fashion - ostia pairs of lateral openings perforating the wall; enable the blood to flow into the heart from the pericardium (large surrounding sinus, a part of the hemocoel) ii. blood - amoebocytes - hemocyanin respiratory pigment - direction of blood flow: heart ===> arteries ==> sinuses (which bathe the tissues) ==> pericardium => heart

VII. Digestive System - foregut derived from the stomodeal region of the ectoderm; chiefly concerned with ingestion, trituration and storage of food; its parts are variously modified for these functions depending upon the diet and mode of feeding - midgut derived from the endoderm; site of en2yme production, digestion and absorption - commonly the surface area is increased by out pocketing forming pouches of large digestive glands - hindgut derived from the proctodeal region of the ectoderm - functions in the absorption of water and the formation of feces VIII. Nervous System A. Brain: - high degree of cephalization accompanied by well-developed sense organs (eyes and antennae). - three major regions: 1. Protocerebrum with one to three pairs of optic centers (neuropiles) which function in integrating photoreception and movement and are probably the centers for the initiation of complex behavior. 2. Deutocerebrum receives the antennal nerves and contains their association centers, arthropods without antennae lack this region. 3. Tritocerebrum gives rise to nerves that innervate the labium (lower lip), the digestive tract (stomatogastric nerves), the chelicerae of chelicerates; commisure is postoral, i.e. located behind the foregut. - with ventral nerve cord and segmental ganglionic swellings B. Sense Organs: - usually associated with some modification of the chitinous exoskeleton which otherwise would at a barrier to the detection of external stimuli. - includes: 1. hairs, bristles, setae that may carry chemoreceptors; when moved, the receptor ending in the shaft or base are stimulated. 2. canals, pits, slits may house chemoreceptors or the opening may be covered by a thin membrane to the underside of which is attached a nerve ending, detect vibrations or other forces that change the tension of the skeleton 3. Proprioreceptors attach to the inside of the integument or to tendons and muscles, for deletion of movement and position of muscles and appendages relative to each other. 4. Eyes a. simple with few photoreceptors b. compound composed of many long cylindrical units called ommatidium - cornea translucent, covers outer part of ommatidium; functions as lens - facet- external surface of cornea is square or hexagonal - crystalline cone long cylindrical or tapered element behind the cornea, functions as a second lens - retinula forms basal end of the ommatidium the center of which is occupied by rhabdome (translucent cylinder) - rhabdomere area of tubules projecting centrally to make up a major part of the rhabdome; functions s a single photoreceptor unit and transmits a signal that represents a single light point - apposition eyes best in bright light - superposition eyes mostly I dim light but may also function in brighter conditions IX. Reproduction and Development - mostly dioecious; - fertilization generally internal in terrestrial forms; external in aquatic forms - eggs rich in yolk, centrolecithal (nucleus surrounded by a small island of non-yolky cytoplasm - cleavage intralecithal or superficial - development stereogastrula

DESCRIPTION OF THE TAXONOMIC GROUPS

SUBPHYLUM TRILOBITOMORPHA (CLASS TRILOBITA)

- fossil trilobites, belonged to the Paleozoic era - Trilobita refers to the transverse trilobation of the dorsal body surface - body oval, flattened with dorsal cuticle; forms are dictated by the mode of existence that may be burrowing, benthic crawling, and planktonic swimming; 3 divisions: a. cephalon fusion ofacron & four post-oral segments; bears a ventral mouth, a pair of ventral antennae, with biramous appendages, distal tips of which bifurcates b. thorax/trunk segments are separated, each bearing a pair of biramous appendages; with telopodite (inner walking leg) & pre-epipodite (outer gill bearing branch) c. pygidium fused segments with reduced appendages; possesses lateral lobes & middle axial lobe arising from 3 longitudinal divisions made by a pair of furrows from anterior to posterior - development 3 larval stages: a. protaspia planktonic with carapace b. meraspia carapace, pygidium & some developing trunk appendages present c. holaspia adult features are evident

SUBPHYLUM CHELICERATA
- body divisions: cephalothorax or prosoma abdomen or opisthosoma - antennae absent - chelicerae. first pair of appendages, feeding structures - pedipalps. second pair of appendages, perform various functions - four pairs of walking appendages

CLASS MEROSTOMATA
- aquatic chelicerates - 5 or 6 pairs of abdominal appendages modified as gills & by a spike-like telson at the end of the body

ORDER XIPHOSURA
- marine soft bottoms, shallow waters - prosoma covered by a large horseshoe-shaped carapace (dorsally convex & ventrally concave) - 2 pairs of eyes: a large dorsolateral & a small dorsomedian pair; both on the carapace - mouth is posterior to the labrum/upper lip (with a pair of trisegmented chelicerae on each side) - 6 pairs of cephalothoracic appendages: a. first 4 pairs identical chelate walking legs - coxae most proximal segment with spines on the median side; macerate & move food anteriorly - flabellum short spatulate process on median side of the last pair's coxae, for cleaning the gills b. fifth pair non-chelate with 4 leaf-like processes; for pushing, clearing, & pushing away the mud & silt during burrowing c. chilaria located between the fifth pair, degenerate & same function of the 5th pair - opisthosoma unsegmented, with 6 short mobile spines at the posterior ends; bears 6 pairs of appendages with the first pair forming the genital operculum - telson a long triangular spine like tail or caudal spine; not a true telson in horseshoe crabs since it does not bear the anal opening; may be used for pushing & for righting the body when it is accidentally turned over - digestion extracellular, intracellular within the hepatic ceaca - excretion & osmoregulation 4 pairs of coxal glands common saclike chambers - coiled tubules ==> bladder ==> excretory pore at the base of the last pair of walking legs - reproduction dioecious with external fertilization; male climbs on the dorsum of the female while the latter scoops out sand to lay from 200 - 300 large eggs which are simultaneously fertilized by the male, after which are covered with sand - development centrolecithal egg ==> cleavage ==> trilobite larva ==> from 13-14 instars ==> adult

ORDER EURYPTERIDA
- extinct giant arthropods that existed from the Cambrian to the Permian period - aquatic, probably attained the largest size of any of the arthropods - smaller cephalothorax - abdomen divided into preabdomen (mesosoma) and postabdomen (metastoma) - six pairs of gills, first pair forms the operculum

CLASS ARACHNIDA
- largest and most important of the chelicerate classes - early arachnids aquatic contemporary living arachnids; terrestrial - migration from aquatic to a terrestrial environment entailed morphological and physiological changes: i. epicuticle became waxy ii. arachnid book lungs and trachea developed from book gills iii. appendages became better adapted for terrestrial locomotion iv. development of silk in spiders, pseudoscorpions; poison glands in scorpions, spiders, and pseudoscorpions - external anatomy body divided into: i. prosoma unsegmented, covered dorsally by a solid carapace; ventral surface covered by coxae of the appendages ii. abdomen segmented, divided into a preabdomen and a postabdomen - appendages arise from the prosoma and consist of a pair of chelicerae (for feeding), a pair of pedipalps, and 4 pairs of legs - respiration by book lungs and/or trachea - circulation heart ==> dorsal anterior and posterior aorta ==> small arteries ==> tissue spaces or sinuses ==> large ventral sinus ==> book lungs ==> venous channels ==> pericardium ==> ostia ==> heart - excretion coxal glands thin-walled spherical sacs along the sides of the prosoma that collect wastes from the surrounding blood; malphighian tubules one or two pairs of slender tubes that arise from the posterior of the mesenteron; wastes pass across its thin syncytial walls and then out into the intestine; nephrocytes large phagocytic cells that are localized in clusters in certain parts of the prosoma and abdomen - nervous system brain: ! protocerebrum contains the optic centers and optic nerves ! tritocerebrum contains the nerves supplying the chelicerae ! resembles a collar or ring surrounding the esophagus since ganglia originally located in the thorax and abdomen have migrated anteriorly and fused with the subesophageal ganglion ! sense organs: i. sensory hairs simple innervated setae trichobothria very long, fine hairs to detect very slight variations or air currents ii. eyes photoreceptors iii. slit sense organs slit-like pits that detect slight changes in the tension of the exoskeleton and also to detect sound vibrations - reproductive system dioecious; single or paired gonads in the abdomen; ducts exit through the genital orifice; sperm transfer is primitive and indirect through spermatophores

ORDER SCORPIONES (Scorpions)

- prosoma covered by a single carapace, long abdomen in a stinging apparatus; chelicerae small, triarticulate, chelate, project anteriorly from the front of the carapace - pedipalps enlarged, form a pair of pincers for capturing prey - abdomen seven segmented preabdomen, five narrow segments of postabdomen - 2 to 5 pairs of small lateral eyes along the anterior lateral margin of the carapace - gas exchange book lungs - excretion 2 pairs of malphigian tubules and a single pair of coxal glands - nervous system distinct nerve cord with 7 unfused ganglia - predaceous - examples Microbuthus pusillus (smallest) Pandinus (largest)

ORDER PSEUDOSCORPIONS (false scorpions)

- lack the long abdomen and sting pedipalp with poison glands - feed on small arthropods such as collembolands and mites - 1 or 2 eyes at each anterior lateral comer or may be absent - gas exchange tracheal system opening through 2 pairs of spiracles on the ventral side of the third and fourth abdominal segments - excretion coxal glands - sense organs indirect eyes, tactile hairs, and trichobothria and lyriform organs - example Chelifer concroides

ORDER SOLIFUGAE (the sun spiders or wind scorpions)

- large arachnids - prosoma large anterior carapace with a pair of closely placed eyes on the anterior median border; short posterior section - abdomen large, broadly joined to the prosoma, segmented - enormous chelicerae projecting in front of the prosoma - pedipalps leg-like but terminate in a specialized adhesive organ used for capturing prey - feed on all types of small animals - gas exchange highly developed tracheal system opening to the outside through 3 pairs of ventral slit-like spiracles on the prosoma and abdomen - excretion pair of coxal glands and a pair of malphigian tubules - example Galeodes graecus

ORDER PALPIGRADI (the palpigrades)

- small; sometimes called microwhip scorpions because of their terminal whip like flagella - carapace is divided into two principal plates between the third and fourth pairs of legs - a tiny third plate is located on each side of the prosoma - pedipalps undifferentiated used as a pair of walking legs - eyes are lacking - abdomen segmented, broadly joined to the prosoma terminates in a long narrow and highly mobile flagellum - feeding habits unknown - some have 3 pairs of booklungs, others reported to lack gas exchange organs - example Koenenia

ORDER UROPYGI (the whip scorpions)

- prosoma covered by a dorsal carapace with a simple pair of anterior eyes, on either side a cluster of 3 to 4 eyes - chelicerae 2 segmented with the distal piece forming a hook or fang - pedipalps stout, heavy, relatively short with the last articles modified to form a pincer for seizing prey - abdomen segmented, the last or 12th segment bearing the anus and a long flagella posterior half with a pair of large anal glands with fluid secretion ( 84% acetic acid and 5% caprylic cid, can burn human skin) - 2 pairs of booklungs on the ventral side of the second and third abdominal segments - excretory organs coxal glands and malphigian tubules - examples Mastigoproctus giganteus, Trithyreus

ORDER SCHIZOMIDA (The Schizomids)

- small tropical arachnids - prosoma divided into 3 segments each covered dorsally by separate plates - abdomen 12 conspicuous segments and bears a terminal flagellum - no eyes - first pair of legs are antenniform - chelicerae composed of 2 pieces - pedipalps leg-like but a little heavier than the true leg - example Schizomus sawadai

ORDER AMBLYPYGI (The Amblypygs)

- tropical and semi-tropical - somewhat flattened body resembling that of spiders - carapace bears a pair of median eyes anteriorly and 2 groups of 3 eyes each laterally - chelicerae similar to those of spiders - pedipalps heavy and raptorial - first pair of legs are modified as long antennae-like appendages with different types of receptors - 2 pairs of book lungs on the ventral side of the second and third abdominal segments - excretory organs a coxal gland and malphigian tubules - example Charinus, Tarantula, Admetus

ORDER ARANEAE (The Spiders)

- tropical mygalomorphs - convex carapace usually with 8 eyes anteriorly, large sternum is present on the ventral surface, and a small median plate known as the labium is attached directly in front of the sternum - chelicerae moderate size; consists of a fang and a basal with a groove into which the fang forms - female pedipalps short and leg-like male pedipalps; modified to become copulatory organs - abdomen globe shape or elongated; segmentation is often reflected in the color pattern; connected to the prosoma by a short and narrow portion called the pedicel - epigastric furrow transverse groove on the anterior ventral side of the body - book lungs with spiracles on each side - spinnerets a group of modified appendages at the end of the abdomen; short conical with many spigots (the openings from the silk glands) - silk of spiders a protein composed of glycine, alanine, serine and tyrosine plays an important role in the life of spiders; used as a dragline and for wrapping up their eggs and the young - predatory feed largely on insects - hunting forms include wolf spiders, fisher spiders, crab spiders, jumping spiders and many mygalomorphs spiders - gas exchange book lungs and/or trachea - circulatory system similar to that of scorpions but the heart varies in length and in the number of ostia - excretion coxal gland is not well developed; malpighian tubules - 8 eyes arranged in 2 rows of 4 each along the anterior dorsal margin of the carapace - example Latrodectus mactans (black widow)

ORDER RICINULEI (The Ricinuleids)

- heavy bodied - anterior margin of the carapace is a hood-like structure that can be raised or lowered - chelicerae 2 segmented and in the form of pincers - pedipalps shorter than the legs, terminate in short pincers not enlarged as those in scorpions - abdomen narrowed in front forming pedicel attach to the prosoma and behind forming a short tubercle that bears the anus; 9 segments are fused and highly sclerotized, coupled to the cephalothorax - feed on other small arthropods - excretory malpighian tubules and a pair of coxal glands - example Ricinoides

ORDER OPILIONES (harvestmen or daddy long legs)

- long legged arachnids - prosoma broadly joined to the short abdomen with no constriction between the 2 divisions - body is rather elliptical in shape - carapace with a tubercle at the center with an eye located on each side - repugnatorial glands along the anterior lateral margins of the carapace; produce secretions often quinones and phenols having an acrid odor; chelicerae small, slender, inarticulate, the last 2 segments forming a pair of pincers; pedipalps short, leg-like similar to those of spiders - abdomen externally segmented; stemites of the first 2 abdominal segments are fused to form an anterior plate known as the operculum which bears the genital openings - predatory, scavenging - excretion pair of coxal glands - respiration through trachea - example Leiobunum rotundum

ORDER ACARI (the mites and ticks)

- lacks body division; abdomen fused with prosoma - capitulum or gnathosoma; head region carrying the mouth parts - rostrum or tectum formed as the dorsal body wall projects - the chelicerae and pedipalps are variable in structure depending on function - chelicerae may become needle-like or terminate in pincers - 4 pairs of legs are each 6 segmented; composed of a coxa, a trochanter, a femur, a genu, a tibia, and a tarsus, the last with a pair of claws - ventral side covered by plates - genital plate bears genital orifice - hairs or setae cover the mite bud - example Tydeus starri

CLASS PYCNOGONIDA (sea spiders)

- marine animals with multiple gonopores, ovigerous legs, the segmented trunk and the additional pairs of walking legs - body narrow, composed of a number of distinct segments; head or cephalon with a ventrally curved cylindrical proboscis at the anterior end; posterior part narrowed to form a short neck bearing on its dorsal surface 4 eyes mounted on a central tubercle - trunk 4 to 6 cylindrical segments with the first one fused with the cephalon; with a pair of large processes projecting laterally from each segment - chelifores compose the appendages homologous to chelicerae of arachnids - chelicerae attached to both sides of the base of the proboscis; relatively short and 1 to 3 segmented, the last segment forming a movable finger - most are bottom dwellers feeding on soft corals, anemones, hydroids, bryozoans and sponges - circulatory system heart or dorsal vessel and a hemocoel - no special organs for gas exchange and excretion - brain beneath the ocular tubercle consists of a protocrebrum and a tritocerebrum - sense organs sensory hairs and fore eyes - example Ascorhynchus castelli

CLASS CRUSTACEA
- includes the only major group of aquatic arthropods - habitat most are marine, but there are many freshwater species and few are terrestrial - structure diverse, unique among arthropods for having 2 pairs of antennae; possess a pair of mandibles and 2 pairs of maxillae; carapace covers all part of the body; appendages are typically biramous - excretory system excretory organs are a pair of blind sacs in the hemocoel of the head that open onto the antennal glands or onto the maxillary gland - respiratory system centers on gills which are typically associated with the appendages - sense organs eyes of 2 types: a. a pair of compound eyes b. a small, median, dorsal, naupliar eye with ocelli - reproduction sexual in the form of copulation - development earliest hatching stage is a naupliar larva which possesses a median naupliar eye and the first 3 pairs of body appendages

SUBCLASS BRANCHIOPODA (brine or fairy shrim p, clam shrim p, water flea)

- small crustaceans with foliaceous trunk appendages - mostly inhabitants of fresh water

SUBCLASS OSTRACODA
- minute crustaceans with body enclosed within a hinged, bivalved carapace commonly impregnated with calcium carbonate - having reduced trunk having no more than 2 pairs of appendages

SUBCLASS COPEPODA (copepods)

- mostly small crustaceans with cylindrical bodies - trunk composed of a thorax bearing 5 pairs of biramous appendages and an abdomen without appendages - first antennae longer than the second pair and held outstretched - compound eyes absent; naupliar eye present

SUBCLASS CIRRIPEDIA (barnacles)

- sessile free-living and parasitic crustaceans with body enclosed with a bivalved carapace - both pairs of antennae reduced or absent

CLASS MALACOSTRACA
- crustaceans with trunk composed of 14 segments which bear appendages - the first 8 compose the thorax and the last 6, the abdomen - female gonopores open onto the ventral side of the 5th thoracic segment and the male onto the 8 - compound eyes present in most species
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ORDER STOMATOPODA ORDER EUPHAUSICEA (krill)

- thoracic appendages biramous and anterior ones not modifed as maxillipeds - gills not tightly enclosed by carapace - examples are marine, pelagic, euphausia

ORDER DECAPODA (decapods)

- first 3 pairs of thoracic appendages are maxillipeds - gills tightly enclosed by sides of carapace

ORDER ISOPODA (isopods)

- dorsoventrally flattened peracaridans with sessile compound eyes and lacking a carapace - thorax with 1 pair of maxillipeds and 7 pairs of pereopods - some of the abdominal segments may be fused with telson

ORDER AMPHIPODA (amphipods)

- laterally compressed peracaridans with sessile compound eyes and lacking a carapace - thorax with 1 pair of maxillipeds and 7 pairs of pereopods - last 3 pairs of abdominal appendages modified as uropods

SUBPHYLUM MANDIBULATA/UNIRAMIA
- terrestrial arthropods with appendages that are primitively unbranched - mandibles are non-jointed, unbranched limbs without palps and are used to handle food from below, not from behind - most distinctive feature single pair of antennae, which are believed to be appendages of the second head somite - have body composed of a head and an elongated trunk with many leg-bearing segments - most require humid environment, because their relatively permeable epicuticle usually lacks the high lipid content found in spiders and insects (lipids more important in repelling water than in reducing water loss) - head bears a pair of antennae and usually ocelli, except in certain centipedes, true compound eyes are never present - mouthparts lie on the ventral side of the head and are directed forward - UPPER LIP and roof of preoral cavity consists of EPISTOME and LABRUM - LOWER LIP formed by either a first or second pair of MAXILLAE - MANDIBLES and HYPOPHARYNX enclosed within the preoral cavity - gas exchange tracheal system in which the spiracles cannot be closed (another path of water loss) - excretion takes place through MALPIGHIAN TUBULES - circulatory system: a. heart dorsal tube extending through the length of the trunk, with a pair of ostia in each segment b. branched system of arteries rarely present - nervous system not concentrated; ventral nerve cord contains a ganglion in each segment - reproductive system indirect sperm transfer by spermatophore is highly developed

CLASS MYRIAPODA
ORDER CHILOPODA
- in temperate and tropical regions mandible bears teeth and a thick fringe of setae, lies beneath the ventro-lateral surface of the head; beneath it, mandibles are two pairs of maxillae - a pair of FORCIPULES covers the mouth appendages - poison claws; appendages of first trunk segment - protection: a. last pair of legs in centipedes b. scolopendromorphs and geophilomorphs with repugnatorial glands on ventral side of each segment c. male lithobiomorph centipedes possess glands on the coxae of the last four pairs of legs (for water uptake and production of pheromones) - locomotion orders of centipedes adapted for running; tergal plates to overcome tendency to undulate geophilomorphs adapted for burrowing - nutrition predacious - gas exchange except in scutegeomorphs, the spiracles of the tracheal system lie in the membranous pleural region above and just behind the coxae - excretion single pair of malpighian tubules - sense organs all geophhilomorphs and some scoloendromorphs lack eyes; other centipedes with few to many ocelli; organs oftomosvary present on the head at the base of the antennae in lithobiomorphs and scutigeromorphs -> for vibration detection and monitoring humidity - reproduction general segment of both sexes carries small appendages called GONOPODS - sperm transmission indirect in centipedes - brood eggs in clusters of fifteen or more - EPIMORPHIC (scolopendromorpha and geophilomorpha)- young display full complement of segments when they hatch - ANAMORPHIC (lithobiomorpha and scutigeromorpha) on hatching, young have only a part of adult complement of segments

SUBORDER GEOPHILOMORPHA - slender centipedes - 31 to 170 pairs of legs - eyes absent - intercalary tergal plates located between tergal plates of more or less equal in length - widely distributed in both temperate and tropical regions - example Geophilus, Strigamia, Mecistocephalus SUBORDER SCOLOPENDROMORPHA - 21 pairs of legs, but some 23 - with or without eyes - tergal plates not alternating in size - distributed throughout the world, especially in the tropics - example Scolopendra, Theatops, Otocrytops

ORDER DIPLOPODA
- millipedes or thousand leggers - secretive; largely-shun light living beneath leaves, soil, logs and bark - presence of diplosegments derived from fusion of 2 originally separate somites; each diplosegment has 2 pairs of legs - head convex dorsally and flattened ventrally; sides are covered by the convex bases of the large mandibles - biting edge of mandible with teeth and a rasping surface - gnatochilarium fused pair of maxillae forming the oral chamber floor - trunk dorsoventrally flattened in flat back millipedes - collum extreme anterior segment, legless, forms a large collar behind the head - integument hard particularly tergites impregnated with calcium salts - surface is smooth - protection protective mechanisms include the: i. calcareous skeleton ii. coiling of the trunk into a spiral when at rest or disturbed iii. repugnatorial glands usually a pair per segment, consist of large secretory sac, components of the secretion include aldehyde, quinones, quinols and hydrogen cyanide - nutrition most are herbivorous, eating mostly on decomposing vegetation; some are omnivorous; digestive tract is typically a straight tube with a long midgut - gas exchange, internal transport and secretion 4 spiracles per diplosegment located on the sterna; each opens in an internal tracheal pouch from which arise different trachea - heart ends blindly at the posterior end of the trunk but anteriorly, a short aorta continues into the head, 2 lateral ostia for each segment - 2 malpighian tubules arise from each side of the midgut - hindgut junction and often long and looped - sense organs eyes may be totally lacking; 2 to 80 ocelli, arranged above the antennae, most are negatively phototactic with integumental photoreceptors; antennae with tactile hairs and peg-like and conelike projections supplied with chemoreceptors - reproduction and development a pair of long Rise tubular ovaries lying between the midgut and ventral nerve cord with 2 oviducts - sperm transfer is indirect - development is anamorphic

ORDER SYMPHYLA
- live in soil and leaf mold - resemble lithobiomorph centipedes - trunk contains 12 leg-bearing segments, caused by 15-24 tergal plates; last 14th segment bears a pair of spinnerets, or cerci, a pair of long, sensory hairs (trichobothria); trunk terminates in a tiny oval telson - head projects in front of laterally placed antennae - mandible covered ventrally by a pair of long, first maxillae, second pair of maxillae are fused, forming a LABIUM - single pair of spiracles opens onto the sides of the head, and the trachea supply only the first 3 trunk segments - structure attached to body wall beneath base of each leg: a. coxal sac - eversible, take up moisture b. stylus - no eyes but 2 organs of Tomosvary are well-developed - genital openings located on the ventral side of 4th trunk segment - eggs laid in clusters of about 8 to 12; parthenogenesis common - development anamorphic, on hatching young have 6 or 7 pairs of legs - example Scutigerella immaculata, Hanseniella

ORDER PAUROPODA
- soft-bodied, rather grub-like animals that inhabit leaf mold and soil - all are minute; abundant in forest litter, widespread in temperate and tropical regions - similar to millipedes in a number of ways: a. first segment (collum) and 11th segment and telson are legless b. dorsal tergal plates are very large and overlap adjacent segments c. 5 of the terger carry a pair of long laterally placed setae *unlike collum of diplopods, pauropod is very inconspicuous dorsally and expanded ventrally - sensory organs on each side of the head is a peculiar/disc-like sensory organ (homologous to organ of Tomosvary of the myriapods) - antennae biramous 1 division terminates in a single flagellum; the other in 2 flagella and a peculiar club shaped sensory structure - mandibles for grinding and piercing lower lip; homologous to gnathochilarium of diplopods, because it apparently represents the first maxillae - most feed on fungi or decomposing plant tissue, but some are predatory - neither heart nor trachea ( except in some primitive species); their absence associated with small size of animals - 3rd trunk segment genital segment - sperm transfer via spermatophore - eggs are laid in humus, either singly or in clusters - development anamorphic, young hatch with only 3 pairs of legs - example Pauropus sylvaticus

CLASS HEXAPODA (INSECTA)

SUBCLASS APTERYGOTA
ORDER THYSANURA (bristle tails, silver fish) ORDER COLLEMBOLA (spring tails)

SUBCLASS PTERYGOTA
- insects are winged or at least derived from winged forms

ORDER EPHEMEROPTERA (mayflies)

- elongated insects with net-veined wings antennae small and mouthparts of short-lived adults vestigial - gradual metamorphosis with aquatic nymphs

ORDER ODONATA (dragonflies and damselflies)

- predacious insects with long, narrow, net-veined wings, large eyes and chewing mouthparts - gradual metamorphosis - nymphs aquatic - dragonflies are stout-bodied

ORDER ORTHOPTERA (grasshoppers, katydids, crickets, roaches, mantids, walking sticks)

- large-headed insects with chewing mouthparts with compound eyes - winged and wingless species - largely herbivorous, at times causing vast crop damage - gradual metamorphosis - femur of hind leg enlarged for jumping in many species

ORDER ISOPTERA (termites, social insects)

- winged and wingless individuals composing the colony - soft-bodied, pale, with abdomen broadly joined to thorax - forewings and hind wings of equal size - gradual metamorphosis

ORDER PLECOPTERA (stone flies)

- adults have long antennae - chewing mouthparts, 2 pairs of well-developed - membranous wings or vestigial wings - abdomen with 2 multi-segmented, caudal cerci of varying length - gradual metamorphosis

ORDER DERMAPTERA (earwigs)

- elongated insects resembling beetles - chewing mouthparts, compound eyes, large forceps like cerci - most have fan-shaped wings and elytra - nocturnal with omnivorous food habits - gradual metamorphosis

ORDER EMBIOPTERA (web spinners)

- small, slender, soft-bodied insects with large heads and eyes - live in silken tunnels and feed on plants - silk glands and spinning hairs located on front tarsi - they are gregarious and many individuals may live together - mostly tropical - gradual metamorphosis

ORDER PSOCOPTERA/CORRODENTIA (booklice, bark lice, and psocids)

- snail, fragile, pale insects with chewing mouthparts - wings membranous and front pair a little larger than hind pair, or wingless - live in a wide variety of habitats-bark, foliage, under stones - some infest buildings and found in books - gradual metamorphosis

ORDER ZORAPTERA
- small, pale, soft-bodied insects like termites with chewing mouthparts - both winged and wingless forms - rare insects living in colonies under dead wood in warm climates - gradual metamorphosis

ORDER MALLOPHAGA (chewing lice and bird lice)

- wingless, flattened insects that live as ectoparasites on birds - eyes re reduced or absent, short legs, small thorax - feed on scales, feathers, hair, skin, dried blood around wounds - gradual metamorphosis

ORDER ANOPLURA
- like chewing lice but mouthparts are adapted for sucking - ectoparasites of birds and mammals - parasitic on domestic animals, head louse and crab louse are parasites of humans - aside from irritation produced is more serious like disease such as typhus fever

ORDER THYSANOPTERA (thrips)

- small, slender insects with mouthparts adapted for rasping and sucking - winged and wingless - wings are narrow with few veins and fringed with hairs - nymph-like early instars but a preadult pupa stage - feed on sap in flowers but some thysanopterans feed on termites and smaller insects

ORDER HEMIPTERA (true bugs)

- piercing and sucking mouthparts - forewings with thickened basal and distal membranous section - hind wings entirely membranous - herbivorous and predaceous - gradual metamorphosis

ORDER HOMOPTERA (cicadas, leafhoppers, aphids)

- herbivorous insects related to the hemipterans - membranous forewings - wings are commonly held in a tent-like position over the body - gradual metamorphosis

ORDER NEUROPTERA (lacewings, ant lions, mantispids, snake flies, dobsonflies)

- adults have chewing mouthparts and long antennae - 2 pairs of similar membranous wings with many veins are held tent-like over abdomen - larvae are predacious and usually terrestrial - complete metamorphosis

ORDER COLEOPTERA (beetles, weevils)

- largest order of insects with hard bodies and chewing mouthparts - adults usually have 2 pairs of wings - most are plant feeders but there are predatory families some are aquatic species - complete metamorphosis

ORDER STRESIPTERA
- minute beetle-like insects - mostly parasitic - only males have wings, for females, forewings reduced to club-shaped appendages - complete metamorphosis

ORDER MECOPTERA (scorpion flies)

- slender-bodied insects, often vividly colored - biting mouthparts prolonged as a beak - most have long narrow wings with many veins - adults are omnivorous and grub-like larvae feed on organic matter - complete metamorphosis

ORDER TRICHOPTERA (caddis flies and water moths)

- soft-bodied insects with 2 pairs of hairy membranous wings and poorly developed chewing mouthparts - larvae are aquatic - complete metamorphosis

ORDER LEPIDOPTERA (butterflies and moths)

- soft-bodied insects with wings, body, and appendages covered with pigmented scales - mouthparts are modified as coiled proboscis for sucking flower nectar - larvae are caterpillars and plant feeders, adults feed little or not at all - complete metamorphosis

ORDER DIPTERA (true flies)

- large orders, all have functional front wings and reduced knoblike hind wings - mouthparts are variable like body form - group includes mosquitoes, horseflies, midges, gnats - adults are often vectors of diseases - larvae usually damage vegetables and domestic animals - complete metamorphosis

ORDER HYMENOPTERA (ants, bees, wasps, sawflies)

- large, varied order, with chewing mouthparts but also modified for lapping and sucking in some forms - winged and wingless species - wings are transparent with few veins - larvae are caterpillars or are grub-like with chewing mouthparts - complete metamorphosis

ORDER SIPHONAPTERA (fleas)

- legs are long with large coxae and are adapted for jumping - have piercing and sucking mouthparts and feed on blood of animals - vectors of bubonic plague - complete metamorphosis

Taxonomic Classification to be used in Zoology 111 PHYLUM ARTHROPODA Subphylum: Trilobithomorpha Class: Trilobita Subphylum: Chelicarata Class: Merostomata Order: Xiphosura (horseshoe crab) Class: Arachnida Order: Scorpiones (true scorpions) Order: Uropygi (whip scorpions) Order: Palpigradi (microwhip scoprpions) Order: Schizomida Order: Amblypygi (tail less whip scorpion) Order: Aranae (spiders) Order: Orthognatha (trapdoor spiders) Order: Labidognatha (7 families) Order: Rinuclei Order: Pseudoscorpiones (false scorpions) Order: Solifugae/Solpugida (wind scorpions) Order: Opilliones (harvestmen or daddy longlegs) Order: Acari (mites, ticks) Class: Pycnogonida (or Pentopoda) sea spiders Subphylum: Mandibulata Class: Myriapoda Order: Chilopoda (centipedes) Order: Diplopoda (millipedes) Order: Symphyla Order: Pauropoda Class: Crustacea Subclass: Malacostraca Order: Isopoda (pillbug, wood lice) Order: Amphipoda (sand fleas) Order: Euphausiacea (krill) Order: Stomapoda (stomatopods) Order: Decapoda (crabs, lobster, shrimps) Subclass: Brachiopoda (brine or fairy shrimp, clam shrimp, water fleas) Subclass: Ostracoda Subclass: Coepepoda Subclass: Pentastomida Subclass: Cirripedia (barnacles) Class: Insecta/Hexapoda

Subclass: Apterygota Order: Thysanura Order: Collembola Subclass: Pterygota Order: Ephemoptera (may flies) Order: Odonata (damselflies and dragonflies) Order: Blattaria (cockroaches) Order: Mantidea (mantids, preying mantids) Order: Isoptera (termites) Order: Grylloblattaria Order: Orthorptera (crickets, katydid, grasshopper, locust) Order: Phasmida (walking stick, walking leaf) Order: Dermaptera (earwigs) Order: Embiidina Order: Plecoptera (stoneflies) Order: Psocoptera (booklice, barklice) Order: Anoplura (sucking lice, crab lice) Order: Mallophaga (chewing lice, biting lice) Order: Thysanoptera Order: Hemiptera (true bugs) Order: Homoptera (aphids, cicads, spittlebugs) Order: Neuroptera (lacewings, dobson flies etc.) Order: Coleoptera (beetles) Order: Strepsiptera Order: Mercoptera (scorpion flies, snow fleas) Order: Siphonoptera (fleas) Order: Diptera (true flies, mosquito) Order: Trichoptera (caddiesflies) Order: Lepidoptera (butterflies, moths) Order: Hymenoptera (wasp, ants, bees)