Introduction to Kinorhyncha and...
Transcript of Introduction to Kinorhyncha and...
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Martin V. Sørensen
Associate Professor and curator of microscopic invertebrates
Natural History Museum of Denmark
Introduction to
Kinorhyncha and Gnathostomulida
Block course on Zoology and Evolution, Biozentrum, Uni. Basel
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Martin V. Sørensen
Associate Professor and curator of microscopic invertebrates
Natural History Museum of Denmark
• Taxonomy, biodiversity, morphology,
and phylogeny
• Kinorhyncha, Priapulida, Loricifera,
Rotifera, Gnathostomulida and
Micrognathozoa
• Metazoan Phylogeny
• Curator of Microscopic Invertebrates
Introduction to
Kinorhyncha and Gnathostomulida
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Edgecombe et al. (2011) Higher-level metazoan relationships: Recent progress and remaining questions. Org. Div. Evol.
Ecdysozoa
Platyzoa
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Platyzoa and Gnathostomulida
Platyhelminthes Gastrotricha Gnathostomulida
Rotifera (Monogononta – Acanthocephala -
Bdelloidea - Seisonidea)
Gnathifera
Micrognathozoa
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Gnathostomulida: Jaw Worms
• Marine, interstitial worms, 100 species
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• Marine, interstitial worms, 100 species
• Ventral mouth leading to blind gut
Gnathostomulida: Jaw Worms
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• Marine, interstitial worms, 100 species
• Ventral mouth leading to blind gut • Simple cephalic sensory organs
• Simple epidermis with monociliated cells
Gnathostomulida: Jaw Worms
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Gnathostomulida: Jaw Worms
• Marine, interstitial worms, 100 species
• Ventral mouth leading to blind gut • Simple cephalic sensory organs
• Simple epidermis with monociliated cells
• Pharynx with hard parts forming jaws
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Gnathostomulida: Jaw Worms
• Marine, interstitial worms, 100 species
• Ventral mouth leading to blind gut • Simple cephalic sensory organs
• Simple epidermis with monociliated cells
• Pharynx with hard parts forming jaws
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Haplognathia lunulifera Gnathostomula paradoxa
Problognathia minima
Mesognatharia remanei
Tenuignathia rikerae
Rastrognathia macrostoma
Gnathostomulida: Jaw Worms
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Two classes: Filospermoidea Bursovaginoidea
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Two classes: Filospermoidea Bursovaginoidea
Reproductive mode: • Hermaphrodite • Soft, muscular penis • Filiform sperm cells Reproductive mode:
• Hermaphrodite • Sclerotized penis stylet and dwarf sperm • Or – non-sclerotized penis with giant sperm cells • Female organs with bursa
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Two classes: Filospermoidea Bursovaginoidea
Order: Scleroperalia
Penis stylet
Testes
Bursa
Ovarium
Spermatozoa
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Gnathostomulida:
Biology and distribution
• Sand mixed with detritus
• Never in mud • Never in clean sand, e.g., high energy
beaches
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Gnathostomulida: Jaw Worms
• Discovered in 1928 by Adolf Remane
• Found again in the 1950’ies and described in 1956 as a turbellarian order (Ax, 1956)
• Raised to class level, but still a Platyhelminthes (Ax, 1964)
• Assigned phylum rank, but with uncertain phylogenetic position (Riedl, 1969)
• Independently, Ahlrichs (1995) and Rieger & Tyler (1995) do comparative studies of gnathostomulids and rotifers
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Gnathostomulida: Jaw Worms
Rotifera Gnathostomulida
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Gnathostomulida: Jaw Worms
Micrognathozoa
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Platyzoa and Gnathostomulida
Platyhelminthes Gastrotricha Gnathostomulida
Rotifera (Monogononta – Acanthocephala -
Bdelloidea - Seisonidea)
Gnathifera
Micrognathozoa
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Kinorhyncha: Mud dragons
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Edgecombe et al. (2011) Higher-level metazoan relationships: Recent progress and remaining questions. Org. Div. Evol.
Ecdysozoa
Platyzoa
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Ecdysozoa
Kinorhyncha (Mud dragons)
Nematoda (Round worms)
Tardigrada (Water bears)
Arthropoda (Arthropods)
Priapulida (Priapulid
worms)
Loricifera (Girdle
wearers)
Nematomorpha (Gordian worms)
Onychophora (Velvet worms)
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Kinorhyncha
• 189 described species
• Marine, meiobenthic
• Present from tidal zone to over 7000 m depth
• Sizes from 130 µm up to 1 mm
• Are composed of a head, a neck and 11 trunk segments
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Kinorhyncha Head
Neck
Trunk
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Kinorhyncha
Mouth cone
Introvert
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Kinorhyncha
Neck
Placids
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Kinorhyncha
Placids
Neck
Placids
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Kinorhyncha
Plate junctions
Segments
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Kinorhyncha
Plate junctions
Segments
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Kinorhyncha
Spines
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Kinorhyncha Development through 6 juvenile stages
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Kinorhyncha: Global Biodiversity
= only few species reported, or kinorhynch fauna known from a single locality only
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Kinorhyncha: Korean Biodiversity Program
• Campyloderes macquariae • Condyloderes megastigma • C. paradoxus • Dracoderes abei • D. orientalis • Dracoderes n. sp. 1 • Echinoderes aspinosus • E. aureus • E. cernunnos • E. koreanus • E. microaperturus • E. obtuspinosus • E. rex • E. tchefouensis • E. ulsanensis • Echinoderes n. sp. 1 • Echinoderes n. sp. 2 • Kinorhynchus yushini • Meristoderes elleae • M. glaber • M. herranzae • M. imugi • Meristoderes n. sp. • Paracentrophyes anurus • Pycnophyes tubuliferus • Pycnophyes n. sp. 1 • Pycnophyes n. sp. 2 • Pycnophyes n. sp. 3 • Pycnophyes n. sp. 4 • Sphenoderes poseidon
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Kinorhyncha: Gulf of Mexico Research Initiative
• Antygomonas n. sp. • Echinoderes n. sp. 1 to 11 • Echinoderes spinifurca • Paracentrophyes n. sp. • Pycnophyes n. sp. • Sphenoderes n. sp.
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Ecdysozoa
Kinorhyncha (Mud dragons)
Nematoda (Round worms)
Tardigrada (Water bears)
Arthropoda (Arthropods)
Priapulida (Priapulid
worms)
Loricifera (Girdle
wearers)
Nematomorpha (Gordian worms)
Onychophora (Velvet worms)
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Ecdysozoa
Kinorhyncha Nematoda Tardigrada Arthropoda Priapulida Loricifera Nematomorpha Onychophora
Ecdysozoa
• Cuticle formed from microvilli tips • Molting • Ecdysone • Loss of locomotory cilia
Scalidophora Nematoida
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Ecdysozoa
Kinorhyncha Nematoda Tardigrada Arthropoda Priapulida Loricifera Nematomorpha Onychophora
Panarthropoda
Segmentation of trunk and limbs
Cycloneuralia
Ring-shaped brain around pharynx
Scalidophora Nematoida
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Cycloneuralia:
Scalidophora and Nematoida
Kinorhyncha Nematoda Priapulida Loricifera Nematomorpha
Scalidophora Nematoida
• Introvert + mouth cone • Scalids • Papillar sense organs (flosculi) • Protonephridia
• Unpaired dorsal and ventral nerve strings • Loss of circular muscles • Loss of sperm flagellum • Both sexes with cloaca • Vermiform habitus
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Molecular data does not support
Scalidophora
Kinorhyncha Nematoda Priapulida Loricifera Nematomorpha
Scalidophora Nematoida
Plesiomorphies? Adult morphology
18S rRNA
• Introvert + mouth cone • Scalids • Papillar sense organs (flosculi) • Protonephridia
• Unpaired dorsal and ventral nerve strings • Loss of circular muscles • Loss of sperm flagellum • Both sexes with cloaca • Vermiform habitus
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Loriciferan – Nematomorph sister groups? Loricifera Nematomorpha
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Loricifera Nematomorpha
• Hexaradial symmetry patterns in pharynx, opposed to pentaradial symmetry patterns in Kinorhyncha and Priapulida
Loriciferan – Nematomorph sister groups?
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Loricifera Nematomorpha
?
Evolution through Progenesis?
Progenesis: Evolution through sexual maturation of larval stage, which causes the loss of following preadult and adults stages
Loriciferan – Nematomorph sister groups?
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Evolution through Progenese
Loss of larva
Original cycle maintained
Loss of adult stage through progenesis
Loriciferan – Nematomorph sister groups?
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The New Ecdysozoan Phylogeny?
Kinorhyncha (Mud dragons)
Nematoda (Round worms)
Tardigrada (Water bears)
Arthropoda (Arthropods)
Priapulida (Priapulid
worms)
Loricifera (Girdle
wearers)
Nematomorpha (Gordian worms)
Onychophora (Velvet worms)
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