Note for BSC 373 with Professor Harris at UA-Lecture 12
Note for BSC 373 with Professor Harris at UA-Lecture 12
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uwm Batrachomorphs vs Reptilomorphs Distinguished in part by the way the roof of the skull is fastened to the posterior portion of brain case Batrachomorphs similar to a hogquot Skull roofattachedto braincase Via the exoccipitals Loss ofskull kinesis Only fourfingers ll l hand Batrachomorphs vs Reptilomorphs Distinguished in part by the way the roof of the skull is fastened to the posterior portion REFTILIOMORPH tropllasic of braincase Batrachomorphs Reptilomorphs similar to a reptilequot Skull roof attached to braincase via the basiocoipital increased skull kinesis exoccrmlals has owp m Batrachomorphs vs Reptilomorphs Batrachomorphs Largest and longest lasting group of primitive nonamniote tetrapods Some lineages of temnospondyls extended into the Cretaceous and at least some living Cums to m amPhibians may be mssunowuoxoa derived from LCquot 4 Penn temnos ond ls x p y 39 l l j l Eryups matm in mmnsnauovu mm LCmL Batrachomorphs vs Reptilomorphs ANTHRACOSAURDlDEA Reptllomorphs Contains a diverse array of animals both nonamniotes and amniotes Early reptilom orphs have a number of skull and jaw modi cations suggesting they fed on terrestrial invertebrates Eniwulunmn quJhc mrldunJvJSm Temnospodylii Temnospondyls are a very large and widespread extinct clade of stegocephalians They are known from the Lower Carboniferous ca 340 mya to the Lower Cretaceous ca 120 mya and have been found on all continents from Greenland to Antarctica More than twelve families are known with about 90 genera Display a great diversity of forms resembling large salamanders crocodiles or gavials with brevirostral shortsnouted to longirostral long snouted skulls Their size ranges from about 20 cm to 3 m in length Temnospodylii Aquatic semiaquatic amphibious or terrestrial this opportunist group occupied a great range of ecological niches especially during the Upper Carboniferous and the Lower Permian From the Upper Permian to their demise in the Lower Cretaceous temnospondyls are largely ifnot entirely restricted to aquatic habitats ie they were found in 39esh waters on continents and in the seas wmvmw m Rayv nned Fish Coelawnms Lung sh Anura 4300 E E Urodela 415 4900 165 Gymnopmona if DJ v39 ParentEmmi A 39LMRIGSquot 3300 U q Am lirsbaenra J r i p E36000 9 S 539 Serpemes2500 2 s g g 2 5 g g Lrlanlsquot T m m Rhynchocephalia Tumara 039 4 777 Testudrnes Chelama 260 lm Lquot quot Crocodylra 24 3 3 1 rA 9000 I Mammalra 4000 ml l r L4 m Lissamphibia The word Lissamphibia contains liss smooth referring to the smooth skin amphi double and bias life referring to the aquatic and terrestrial life stages that characterize many amphibians though not all Three extant lineages Anura Salienta frogs and toads north to south temperate Urodela Caudata salamanders north to south temperate in Americas subtropical to north temperate in Eurasia Gymnophiona Apoda caecilians tropical and subtropical Earliest Lissamphibian fossil is from early Triassic ca 240 mya gtTriadobatrachus an anuran Earlierfossil material exists Permian that may represent amphibians Salamanders and caecilians present in early Jurassic ca 200 mya Lissamphibia Super cially each of the extant lineages are very different from each other Possibly evolved from a salamanderlike ancestor Differences between lineages related to locomotor specialization Adults of all living species are carnivorous Opportunistic feeders Only larval anurans tadpoles are herbivorous Lissamphibia Synapomorphies Pedicellate bicuspid teeth the crown and base or pedicel of teeth composed of dentine separated by a zone of uncalcified dentine made of fibrous connective tissue This feature is found in living amphibians and some temnospondyls pedicellate teeth 39 crown 1 pe Ice Pough et al 2001 Herpetology Ii I entice Hall Inc A39A d inner surfaceo Jaw pleu rodont attachment Lissamphibia Synapomorphies Stapes columella and operculum are sound conducting structures The stapes is known only in tetrapods The operculum is connected via opercular muscle to the suprascapula Caecilians lack opercular muscles no girdle There are two sensory patches in the inner ear the papilla basilis is found in other tetrapods and the papilla amphibiorum is unique to living amphibians The stapes is directed dorsolaterally from fenestra ovalis Two occipital condyles on skull and two cotyles on atlas Reduction in skull elements loss of supratemporals intertemporals tabular postparietals jugals postorbitals outer inner ear middle ear le arl lei papilla amphibiorum a 4 gt1000 HZ Wailquot borne waves 1quot 394 stapes L tympanum K i 9 AMA stapedial m 1 papilla el basiiarisir operculum suprascapular 391 OPeFCU39ar mv opercularis m operculum seismic ground borne waves lt 1 000 Hz Anuran Auditory System Duellman amp Trueb 1994 Biology of Amphibians HEARING IN AMPHIBIANS squamosal seismic sound waves Urodele Auditory System lunzey 2001 Vertebrate Biology Lissamphibia Synapomorphies vertical canals 900to each other horizontal canal papilla amphibiorum tr39culus 1001000Hz papilla basilaris sacculus 10002000HZ sensory patChes lagena amphibians otoliths cochlear duct reptiles after Porter 1972 Herpetology W B Saunders 00 Toronto Lissamphibia Synapomorphies paired occipital condyles stapes directed dorsolaterally Lissamphibia Synapomorphies 39 Levator bulbi muscle that elevates the eye this is a thin muscle in the floor of the orbit innervated by the 5th cranial nerve that causes the eye to bulge outward and to enlarge the buccal cavity This muscle is present in anurans and urodeles and in a modified form in caecilians 39 The fat bodies develop from germinal ridge synapomorphy 39 The skin contains mucous and granular poison glands 39 Cutaneous respiration all amphibians rely on the skin surface for gas exchange especially for the release of 002 Amphibians show increasing reliance on the lungs for 02 uptake as temperature and activity increase St 9 E L D 399 Q LLI Mucous Gland With Duct Granular Gland Chromatopho Rana lessonagz ratum corneum res 4 AMPHIBIAN SKIN Pattamp Patt 1969 Comparative Vertebrate Histology Harperamp Row Publ New York xanthophores pterydine carotenmds m I red yellow orange WRanaggv hs 39 iridiphores purine granules white Silver melanophores 7 melanin black brown dendritic pocesses of melanophores after Pough et al 2001 Herpetology PrenticeHall Inc Lissamphibia Synapomorphies 39 Short straight ribs 39 Green rods in frog and salamander retinas In vertebrate retinas with dual vision there are two classes of photoreceptors cones that are used in bright light conditions during the day and rods that are used in dim light They both consist of an inner segment where metabolic processes take place and an outer segment that is packed with visual pigment molecules Vision is initiated in the outer segment Green vs red rods associated with terrestrial vs aquatic habitats Structure of a Frog Retina Green rod 432 nm Red rod 502 nm pigment epithelium receptor segments Sin le Cone 80 al 8 W Double Cone Accessory Princip after Gordon J 8 I C Hood 1976 Anatomy and pr SiOlOgV of thefrog retina IN The Amphibian Visual System A MLI licliscrplinary Approach K V Fits ed Academic Press in New York 2986 Lissamphibia Synapomorphies Short straight ribs Green rods in frog and salamander re inas They are the most pri tive group of vertebrates to have a 3chambered heart and double circulation Anurans Frogs toads and their close fossil relatives Anura means without a tailquot Frogs outnumber salamanders and caecilians substantially There are 165 living species ofcaecilians 415 living salamanders and ca 4300 species offrogs Anuran Synapomorphies I Frogs have at most nine vertebrae in front ofthe sacmm and the three or four posterior to the sacrum are fused into a rod called the urostyle In contrast caecilians and salamanders have many more vertebrae and they do not have a urostyle Frogs do not possess tails in the adult stage as caecilians and salamanders do Anuran Synapomorphies I Frogs also have a radioulna a fused radius and ulna bones ofthe forearm and a tibiofibula fused tibia and bula bones of the shank The tibiale and fibulare ankle bones aka astragalus and calcaneum of 39ogs are greatly elongate results in an additional lever system hat frogs can utilize in jumping Origin of saltmion and its morphological correlates lack ofa tail reduction in vertebrae elongation of propulsive segments of the body seems to be one of the features that clearly sets li39ogs apart li39om other major vertebrate groups Anurans Distinctive life history phase known as the tadpole which is a highly specialized quoteating machinequot Salamanders and caecilians have a larval form but in neither does the larva possess the many specializations such as the ceratohyal pump that 39og tadpoles have Even the most basal living li39ogs have the beginnings of a unique mechanism of tongue projection that is associated with extreme modification ofthe gill arches into a fused hyobranchial plate Anurans Frogs vs toads Not natural groupings rather distinction is based on morphological and life history characters Frogs Relatively longer hindlimbs for longerjumps Aquatic to semiaquatic Well developed webbing on feet Relatively smooth skin Toads More stout body with shorter hindlimbs Less aquatic webbing reduced or absent Skins exhibit a rougher texture Anurans Other phenotypes Arboreal Large heads eyes may be directed forward Slim waists and long thin limbs Walking and climbing as well asjumplng Enlarged adhesive toe disks may be present Strictly aquatic ie pipids Dorsoventrally flattened Thick waists and powerful hindlimbs extensive webbing Lateral line system Do not have a tongue Anuran Diversity Megophryidae They are found in India Pakistan and eastward into southeast Asia Borneo and the Philippines to the Sunda lslands They range in size from about 20 to 125 mm Their skin is modified so that they resemble dead leaves on the forest floor Some species have points of skin on the eyelids which further enhances the illusion Megophrys montana 10 Anurans Rhinophrynidae The Mexican Burrowing Toad Rhinophrynus dorsalis is the only living member ofthis family Highly fossorial adapted for burrowing and apparently only comes to the surface to breed and then only during very heavy rains The head is small and coneshaped and not surprisingly the skull resembles that of a mole The skin is quite thick and an early naturalist described the frog in French as a bag of bones edge of the foot the first toe is also modified to extend the functional edge of the spade Rhinophrynus is specialized for eating termites and the tongue seems to be particularly modified for this activity Instead of being flipped out over itself as in most frogs the tongue is projected straight out of the mouth There is a spade used for digging on the inner a quotW Anuran Diversity Pipidae Tongueless Frogs Aglossa Highly aquatic frogs that rarely if ever venture out of water Several adaptations to aquatic life loss of the tongue not generally useful for feeding in water presence of lateral line organs Surinam Toad Pipa pipa is extremely flattened and looks like a road kill Females of Pipa have an elaborate mating behavior eggs are deposited on the back ofthe female and the skin swells up around the eggs to encase them in pockets in which the embryos develop In some species the eggs hatch out as tadpoles but in others fully formed froglets emerge from the mother39s back Anuran Diversity Alla miyne rulhvenl Brah e nahaae Triauubalvachus massinun e W 7Vieraella nemsn Helen m ne 7 Nolobalraclius degmsloi e Leglndarlylldaa Liinnac llaSll ae Ascaghus tillel Leiogelma 7Eodiscoglossussanionaef G M ODHUECWME B Bumbmalundae Soonlassmae g s C Disco in sidae Rhmodemd Eu elcbalinae C Megcghryidae H age Pelcbalidae H W F Peludylidae Ceiilralemdae Rhinughrymdae Mmquot as 4 H pig s Pipidae W Necbaracma 0 A mlhiole ISIS Ranidae Hy El llldae Rn acognandae 11 Anuran Diversity Rhinoderma The genus is known from the temperate forests of southern Chile and Argentina The frogs are small about 30 mm long In Rhinoderrna danvinii the males carry the tadpoles in their vocal sacs and development is completed as froglets Anuran Diversity Hylidae Treefrogs One of the largest families of frogs about 3739 genera These are arranged in four subfamilies the first three of which are distinctive Pelodryadinae Phyllomedusinae Hemiphractinae and Hylinae Pelodryadinae Australian treefrogs completely terrestrial and lack the expanded toepads that are typical of most hylids Anuran Diversity Hylidae Phyllomedusinae Leaf Frogs occurs from Mexico through tropical South America Redeye tree frog is most famous species Leaf frogs generally deposit their eggs on vegetation near ponds and the hatchling tadpoles fall into the water one genus has highly streamadapted tadpoles with funnelmouths 12 Anuran Diversity Hylidae Hemiphractinae these frogs care for the developing eggs by carrying them on their back in a shallow basin or in pouches modified from the skin The developing embryos hatch as froglets in some species or as tadpoles in others Anuran Diversity Hylidae Hylinae remainder of the tree frogs are placed in the Hylinae because they do not possess the distinctive features of the other subfamilies Among the most bizarre hyline frogs are certain casque headed genera such as Triprion and Trachycephaus in which the skull bones are elaborated into a solid helmet Certain casque headed species use their bony heads to block the entrances to their burrows and reduce evaporative water loss Anuran Diversity Dendrobatidae Poisondart frogs As a group they are the most brightly colored of frogs They are also mostly diurnal and observed hopping on the forest oor by day Diverse reproductive behaviors g4 For groups with data the tadpoles are carried on the back ofthe adult in some species it is the male in 39 others it is the female that carries the tadpoles Tadpoles are transported to a body of water usually a stream but also small ponds the waterfilled axils of bromeliads or some other small container ln Dendrobates females transport one tadpole at a time and there is only one tadpole per hiding place These hiding places offer little in the way of food 39 resources to the developing lava and the female vs samu has evolved the remarkable behavior of depositing unfertilized eggs in the axil to feed the developing tadpole iLlntfvrahll Raymtg Ftsh A Arm cs 5 Luams 33m Amputathva serpmnyszsm umdwwi L ms Rhynmncemlha Ttmliru mm chmm 25b Urodela Satarnanoers are orstrngursneotrorn otnerarnpnrbrans trogs and oaecmans rnostobyrousty by ne presence of ataH tn 5 taryae tuyennes and aoutts and by nayrng hmbs set at ngnt angtes to ne body wrtn forehmbs and mndhmbs of approxtmate y equat srze except m tne famHy Strenroaewnrcn tacks nrnonrnbs Urodela Synapomorphies Satarnanders are drstrngursned from frogs and caecmans atso by numerous cnaractenstrcs of tnerr sketeton and muscu ature 7 Presence or a tart and usuaHy two parrs or hmbs or approxrrnatety equat Sm 7 Absence or skuH bones pusiurbttak pus tpartetaL tabu ar supratempuramugak euadratcyugat present rn Karaurtdae supraucmpttah basroccrprtat and Ectupterygmd bones 7 Presence uftrue teetn un butmavvs 7 3m SM and externat grtts rn aquatu tawae when present 7 Absence or an otrc notcn and rnreete ear 14 Urodela Many salamanders have a biphasic life cycle containing an aquatic larval form with external gills and a metamorphosed terrestrial adult form that breathes by lungs andor through moist skin Some species lack metamorphosis and retain a larval appearance throughout their life whereas other species lack the aquatic larval stage and hatch on land as terrestrial forms that resemble miniature adults The general association of lungs with terrestrial animals and gills with aquatic ones is not universally true among salamanders Terrestrial plethodontid salamanders lack lungs and breathe through the skin whereas aquatic adult amphiumas lack gills and use lungs to breathe Salamanders are carnivorous in the larval juvenile and adult stages oftheir life cycles they prey mainly on small arthropods Marbled salamander Urodela world where moist and cool habitats occur Salamanders inhabit nearly all northern temperate regions of the The greatest phylogenetic diversity of salamanders is in North America where representatives of 9 of the 10 extant taxonomic families occur ca 420 species Lungless salamanders ofthe family Plethodontidae occur also in tropical areas of Central and South America a few species also occur in southern Europe rocks or logs from which they emerge at night to feed stones Terrestrial salamanders are usually found on the forest floor under Aquatic forms live on the bottoms of streams or ponds often under Regardless of habitat terrestrial or aquatic all are aquatic breeders Salamanders are usually seasonal in their activity and are easily found only during their active season which differs among species Urodela Paedomorphosis is common Larval or larvallike form becomes sexually mature without attaining adult body form Retention of larval characters larval teeth bone patternsstructure absence of eyelids lateral line system external gills gill slits Most common in aquatic forms In some salamander species individuals either transform from an aquatic larval stage into a terrestrial quotmetamorphic adultquot or they remain within the aquatic environment as a sexually mature quotpaedomorphic adultquot 39 This dimorphism termed facultative paedomorphosis is environmentally induced because environmental conditions experienced as a larva in uence whether an individual becomes one morph or the other Life Cycle of Facullah ve Paedomorphs 15 Urodela Most species exhibit internal fertilization via spermatophores pyramid shaped plugs of sperm Sperm cap on a gelatinous base Only two families have external fertilization Essentially all species are oviparous A few soecies of salamandrids are vivioarous U rodela Amphiumidae Plelnodomidae Salamandridae Dicamgtodontidae Ambystomatidae Proteidae Rhyacotritonidae Cryptobranchidae Hynobiidae Sirenidae Batrachosauaroididae f Karauridae v Prosirenidae f 2 Scapherpetontidae t Amphiumidae Elongate length may exceed 1 meter and permanently aquatic although adults lack gills and use lungs to breathe air Both forelimbs and hindlimbs are extremely reduced in size and appear nonfunctional although the animals move them Three extant species of the genus Amphiuma differ in the number of toes present on he limbs from one to three Larvae hatch with external gills and transform partially losing the gills but retaining one pair ofgill slits 16 Amphiumidae Amphiumas inhabit swamps bayous marshes drainage ditches and streams in the southeastern United States Amphiumas are nocturnally active generalist predators that feed on crayfish frogs fish and small snakes They serve as prey for larger snakes including cottonmouths and mudsnakes Large amphiumas have a powerful bite that is unusual for salamanders and they should not be handled carelessly Breeding has been observed for the twotoed and three toed amphiumas These amphiumas breed in the winter or spring fertilize eggs internally and lay a string of eggs in muddy shallow water Eggs are guarded un il they hatch in the summer or fall Plethodontidae Lungless salamanders are widely distributed in eastern and western North America and Central America with populations also in South America southern Europe and Sardinia The family Plethodontidae is by far the largest salamander family wi h approximately 250 recognized species All plethodon id salamanders are lungless and breathe through moist skin Plethodontidae Plethodon ids are unique among salamanders in having narrow grooves between each nostril and the upper lip Males often have protuberances on the upper lip associated with the nasolabial grooves and a mental gland located beneath the mouth Costal grooves are pronounced Most plethodontids are completely terrestrial and lay eggs on land Some have a biphasic life cycle with an aquatic larva and terrestrial adult and others are completely aquatic and permanently larval in form Aquatic forms usually inhabit streams 17 Salamandridae o Salamanders of the family Salamandridae are informally subdivided into two major subgroups the quottrue salamandersquot includes genera Chiogossa Mertensiela and Salamandra and the newts o Both groups have aquatic larvae except for some viviparous true salamanders Mertensiela luschani Salamandra atra and Salamandra salamandra bernardezi that give birth to fully metamorphosed offspring Metamorphosed adults of the true salamanders are highly terrestrial whereas those of newts are at least partly aquatic and some are entirely aquatic Salamandridae o Some newts of the North American genus Notophthamus have a particularly complex life cycle with two metamorphoses and three distinct developmental stages an aquatic larva a terrestrial juvenile the quotred eftquot and a secondarily aquatic adult o True salamanders are Eurasian in distribution newts are found in Eurasia Africa N America o All salamandrids have toxic skin secretions and newts are highly poisonous in all stages of their life history Many salamandrids have bright colors that serve as warnings of their toxicity and may be used in defensive displays Salamandridae o Newts of the family Salamandridae are unlike all other salamanders in having roughtextured skin that is not slimy o Costal grooves usually are not distinct o Most salamandrids have a biphasic life history with aquatic larvae and metamorphosed adults that may be terrestrial or aquatic The tail is finlike in aquatic forms but not in terrestrial forms Larvae have four pairs of gill slits and large external gills Legs are relatively long prominent and robust 18 Sexual Dimorphism in Newts Proteidae Mudpuppies or waterdogs Neoturus Salamanders of the family Proteidae are permanently aquatic and maintain throughout life the large filamentous gills and caudal fins characteristic of larvae r r Cryptobranchidae Hellbenders Elongate length may exceed 15 meters in the genus Andrias dorsoventrally flattened and permanently aquatic The genus Andrias contains the largest living salamanders and fossil species attained even larger sizes 1 exceeding 2 meters in body length Fleshy dermal folds are visible along the lower sides of the body in adults Larvae have caudal fins and short external gills Metamorphosis is partial external gills quot are lost in adults but eyelids do not develop and he tail remains laterally compressed and finlike They live under rocks on the bottoms of fast flowing streams and rivers 19 Sirenidae Sirens do not metamorphose and retain the appearance of an aquatic larva throughout life They are long and slender often described as eellike in appearance with external gills gill slits small forelimbs and no hindlimbs The complete absence of hindlimbs distinguishes sirens from all other salamanders Sirenidae Sirens are often nocturnal in their activities and spend the day under cover objects burrowing in mud and weeds Glands in the skin can create a moistureretaining seal overthe body permitting sirens to aestivate in mud burrows when the shallow water of their habitats evaporates Gymnophiona Caecilians Caecilians are amphibians that resemble large earthworms in shape but are a different color yellow or even purple Their mouths however are large and they are predacious upon various small sh or invertebrates They tend to burrow in both the wild and in captivity Found throughout the world39s tropical regions comprised of 6 families which are represented by approximately 35 genera and ca 165 species Most caecilians are fossorial and remain underground throughout most of their lives while other forms are aquatic 20 Gymnophiona Caecilians Oviparous or Viviparous After the developing embryos use up the last oftheir yolk they still require nourishment while inside their mother39 s oviduct The embryos begin feeding upon a substance labeled uterine milk which is secreted by the oviducts The embryos are equipped with tiny uterine teeth that may be used to scrape away and consume the oviduct secretion The teeth are then lost before or shortly after birth Gymnophiona Caecilians Synapomorphies Caecilians are limbless and completely lack pectoral and pelvic girdles Many species have eyes completely or partially covered by bone or esh A tentacle is present on either side ofa caecilian39 s head between the eye and the nares These tentacles are apparently signi cant in receiving chemosensory cues Eversible male copulatory organ or phallodeum that is partially formed by the cloacal wall and by which internal fertilization is accomplished An absence of ear drums or inner ear cavities The means by which the jaws operate via dualclosing musculature is an obvious adaptation to facilitate a burrowing lifestyle Gymnophiona Caecilians Synapomorphies Annuli rings are found throughout the body of most caecilians Caecilians are the only amphibians with scales These dermal scales are located within the annuli and require very close inspection and the use ofa good microscope to notice 21
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