464:
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179:
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623:. The small, long, thin pycnogonid heart beats vigorously at 90 to 180 beats per minute, creating substantial blood pressure. The beating of the sea spider heart drives circulation in the trunk and in the part of the legs closest to the trunk, but is not important for the circulation in the rest of the legs. Hemolymph circulation in the legs is mostly driven by the
869:, which was taken as evidence that Pycnogonida may be a sister group to all other living arthropods, the latter having evolved from some ancestor that had lost the protocerebral appendages. If this result had been confirmed, it would have meant the sea spiders are the last surviving (and highly modified) members of an ancient stem group of arthropods that lived in
2716:
684:. The abdomen and the thorax with its thoracic appendages develop later. One theory is that this reflects how a common ancestor of all arthropods evolved; starting its life as a small animal with a pair of appendages used for feeding and two pairs used for locomotion, while new segments and segmental appendages were gradually added as it was growing.
584:
is the only family where both the chelifores and the palps are functional. In the others the chelifores or palps, or both, are reduced or absent. In some families, also the ovigers can be reduced or missing in females, but are always present in males. In those species that lack chelifores and palps,
687:
At least four types of larvae have been described: the typical protonymphon larva, the encysted larva, the atypical protonymphon larva, and the attaching larva. The typical protonymphon larva is most common, is free living and gradually turns into an adult. The encysted larva is a parasite that
2432:
Ballesteros, Jesús A; Setton, Emily V W; Santibáñez-López, Carlos E; Arango, Claudia P; Brenneis, Georg; Brix, Saskia; Corbett, Kevin F; Cano-Sánchez, Esperanza; Dandouch, Merai; Dilly, Geoffrey F; Eleaume, Marc P; Gainett, Guilherme; Gallut, Cyril; McAtee, Sean; McIntyre, Lauren (2021-01-23).
849:
A competing hypothesis proposes that
Pycnogonida belong to their own lineage, distinct from chelicerates, crustaceans, myriapods, or insects. This Cormogonida hypothesis contended that the sea spider's chelifores, which are unique among extant arthropods, are not positionally homologous to the
2371:
Ballesteros, Jesús A; Santibáñez-López, Carlos E; Baker, Caitlin M; Benavides, Ligia R; Cunha, Tauana J; Gainett, Guilherme; Ontano, Andrew Z; Setton, Emily V W; Arango, Claudia P; Gavish-Regev, Efrat; Harvey, Mark S; Wheeler, Ward C; Hormiga, Gustavo; Giribet, Gonzalo; Sharma, Prashant P
416:, found in oceans around the world. The over 1,300 known species have leg spans ranging from 1 mm (0.04 in) to over 70 cm (2.3 ft). Most are toward the smaller end of this range in relatively shallow depths; however, they can grow to be quite large in
877:
expression patterns demonstrated the developmental homology between chelicerates and chelifores, with chelifores innervated by a deuterocerebrum that has been rotated forwards; thus, the protocerebral Great
Appendage clade does not include the Pycnogonida.
1605:
Regier, Jerome C.; Shultz, Jeffrey W.; Zwick, Andreas; Hussey, April; Ball, Bernard; Wetzer, Regina; Martin, Joel W.; Cunningham, Clifford W. (2010). "Arthropod relationships revealed by phylogenomic analysis of nuclear protein-coding sequences".
749:, to the north and south poles. They are most common in shallow waters, but can be found as deep as 7,000 metres (23,000 ft), and live in both marine and estuarine habitats. Pycnogonids are well camouflaged beneath the rocks and among the
862:
and found in the first head segment that in all other arthropods give rise to the eyes and the labrum. This condition of having protocerebral appendages is not found anywhere else among arthropods, except in fossil forms like
2635:
Here we report the first known occurrence of fossil pycnogonids from rocks of
Ordovician age, bridging a 65 Myr gap between controversial late Cambrian larval forms and a single documented Silurian specimen. The new taxon,
1850:
Bastide, A.; Peretti, D.; Knight, J. R.; Grosso, S.; Spriggs, R. V.; Pichon, X.; Sbarrato, T.; Roobol, A.; Roobol, J.; Vito, D.; Bushell, M.; von Der Haar, T.; Smales, C. M.; Mallucci, G. R.; Willis, A. E. (2017).
608:, though sometimes the eyes can be missing, especially among species living in the deep oceans. The abdomen does not have any appendages, and in most species it is reduced and almost vestigial. The organs of this
2057:
Jager, Muriel; Murienne, Jérôme; Clabaut, Céline; Deutsch, Jean; Guyader, Hervé Le; Manuel, Michaël (2006). "Homology of arthropod anterior appendages revealed by Hox gene expression in a sea spider".
897:), though few other relationships at the base of the chelicerates are well resolved. The first phylogenomic study of sea spiders was able to establish a backbone tree for the group and showed that
615:
The morphology of the sea spider creates an efficient surface area-to-volume ratio for respiration to occur through direct diffusion. Oxygen is absorbed by the legs and is transported via the
803:
The class
Pycnogonida comprises over 1,300 species, which are normally split into eighty-six genera. The correct taxonomy within the group is uncertain, and it appears that no agreed list of
1366:). Some of these specimens are significant in that they possess a longer 'trunk' behind the abdomen and in two fossils the body ends in a tail, something never seen in living sea spiders.
1575:
688:
hatches from the egg and finds a host in the shape of a polyp colony where it burrows into and turns into a cyst, and will not leave the host before it has turned into a young juvenile.
691:
Little is known about the development of the atypical protonymphon larva. The adults are free living, while the larvae and the juveniles are living on or inside temporary hosts such as
3628:
560:. The anterior region (cephalon) consists of the proboscis, which has fairly limited dorsoventral and lateral movement, the ocular tubercle with eyes, and up to four pairs of
2006:
Maxmen, Amy; Browne, William E.; Martindale, Mark Q.; Giribet, Gonzalo (2005). "Neuroanatomy of sea spiders implies an appendicular origin of the protocerebral segment".
703:
legs of the father, where it will stay until it has turned into a small and young juvenile with two or three pairs of walking legs ready for a free-living existence.
3479:
3234:
3518:
3260:
2216:
Sharma, Prashant P.; Kaluziak, Stefan T.; Pérez-Porro, Alicia R.; González, Vanessa L.; Hormiga, Gustavo; Wheeler, Ward C.; Giribet, Gonzalo (November 2014).
2734:
3623:
1316:), though some researchers have argued that this putative larval sea spider is not a pycnogonid at all. Unambiguous sea spider body fossils include the
589:
is well developed and flexible, often equipped with numerous sensory bristles and strong rasping ridges around the mouth. The last segment includes the
3453:
3208:
1579:
756:
Sea spiders either walk along the bottom with their stilt-like legs or swim just above it using an umbrella pulsing motion. Sea spiders are mostly
485:
include species with both five and six pairs. There are nine polymerous (i.e., extra-legged) species: seven species distributed among four genera (
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3247:
1583:
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to the rest of the body. The most recent research seems to indicate that waste leaves the body through the digestive tract or is lost during a
1990:
1750:
2785:
631:
as well as a nervous system consisting of a brain which is connected to two ventral nerve cords, which in turn connect to specific nerves.
1952:
604:
make up the extremely reduced body of the pycnogonid, which has up to two pairs of dorsally located simple eyes on its non-calcareous
473:
Sea spiders have long legs in contrast to a small body size. The number of walking legs is usually eight (four pairs), but the family
2316:"Ordered phylogenomic subsampling enables diagnosis of systematic errors in the placement of the enigmatic arachnid order Palpigradi"
680:
The larva has a blind gut and the body consists of a head and its three pairs of cephalic appendages only: the chelifores, palps and
2134:"The chelifores of sea spiders (Arthropoda, Pycnogonida) are the appendages of the deutocerebral segment: Chelifores of sea spiders"
1709:
3590:
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252:
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2314:
Ballesteros, Jesús A.; Santibáñez López, Carlos E.; Kováč, Ľubomír; Gavish-Regev, Efrat; Sharma, Prashant P. (2019-12-18).
3505:
1782:
Woods, H. Arthur; Lane, Steven J.; Shishido, Caitlin; Tobalske, Bret W.; Arango, Claudia P.; Moran, Amy L. (2017-07-10).
1380:
463:
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3143:
3130:
2267:"A Critical Appraisal of the Placement of Xiphosura (Chelicerata) with Account of Known Sources of Phylogenetic Error"
1657:
Sharma, P. P.; Kaluziak, S. T.; Perez-Porro, A.R.; Gonzalez, V. L.; Hormiga, G.; Wheeler, W. C.; Giribet, G. (2014).
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in the part of the gut that extends into every leg, a process called gut peristalsis. These creatures possess an
413:
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1320:
2720:
2374:"Comprehensive Species Sampling and Sophisticated Algorithmic Approaches Refute the Monophyly of Arachnida"
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699:. When the attaching larva hatches it still looks like an embryo, and immediately attaches itself to the
3554:
3393:
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3148:
2217:
2652:. It is the only record thus far of a fossil sea spider in rocks of demonstrably shallow marine origin.
2133:
1110:
1080:
3572:
1908:
Bain, B. A. (2003). "Larval types and a summary of postembryonic development within the pycnogonids".
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2015:
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946:
2435:"Phylogenomic Resolution of Sea Spider Diversification through Integration of Multiple Data Classes"
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Phylogenomic
Resolution of Sea Spider Diversification through Integration of Multiple Data Classes
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extend throughout many appendages because its body is too small to accommodate all of them alone.
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352:
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2266:
2218:"Phylogenomic Interrogation of Arachnida Reveals Systemic Conflicts in Phylogenetic Signal"
1659:"Phylogenomic Interrogation of Arachnida Reveals Systemic Conflicts in Phylogenetic Signal"
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2019:
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791:, which are much larger, most sea anemones survive this ordeal, making the sea spider a
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1979:
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1135:
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chelicerae of
Chelicerata, as was previously supposed. Instead of developing from the
3612:
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2202:
2149:
1974:
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851:
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1937:
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1402:
discovered about 70 fossils from three distinct species in the 160-million-year-old
854:
like the chelicerae, the sea spider chelifores were thought to be innervated by the
451:, if correct. This is disputed, however, as genetic evidence suggests they may be a
3419:
3135:
2997:
2972:
2962:
2933:
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2165:
2094:
2043:
1853:"RTN3 is a Novel Cold-Induced Protein and Mediates Neuroprotective Effects of RBM3"
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1302:
The fossil record of pycnogonids is scant. The earliest fossils are known from the
1189:
1073:
865:
711:
511:
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436:
141:
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1929:
576:, which are used for cleaning themselves and caring for eggs and young as well as
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3174:
3102:
2977:
2891:
2842:
2590:
Rudkin, Dave; Cuggy, Michael B.; Young, Graham A.; Thompson, Deborah P. (2013).
1733:
Crooker, Allen (2008). "Sea
Spiders (Pycnogonida)". In Capinera, John L. (ed.).
1414:. The find will help fill in an enormous gap in the history of these creatures.
1102:
1062:
Hoek, 1881 (Synonyms: Pycnogonoidea Pocock, 1904; Rhynchothoracoidea Fry, 1978)
931:
882:
881:
Phylogenomic studies place the
Pycnogonida as the sister group to the remaining
815:
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624:
609:
605:
581:
517:
505:
482:
220:
165:
149:
112:
45:
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525:. Instead, gasses are absorbed by the legs and transferred through the body by
3025:
2967:
2943:
2826:
2684:
2592:"An Ordovician Pycnogonid (Sea Spider) with Serially Subdivided 'Head' Region"
1876:
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2814:
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2649:
2283:
2265:
Ballesteros, Jesús A; Sharma, Prashant P (2019-11-01). Halanych, Ken (ed.).
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95:
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2086:
2035:
1894:
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located dorsally in relation to the digestive tract. Reproduction involves
650:
All pycnogonid species have separate sexes, except for one species that is
17:
2743:
2715:
1704:. Barnes, Robert D. (6th ed.). Fort Worth, TX: Saunders College Pub.
1390:
In 2007, remarkably well preserved fossils were exposed in fossil beds at
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50:
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2179:
Dunlop, J. A.; Arango, C. P. (2005). "Pycnogonid affinities: A review".
2109:"Chelifores, chelicerae, and invertebrate evolution | ScienceBlogs"
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1307:
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553:
444:
428:
200:
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2640:
n. gen. n. sp., from the Upper
Ordovician (ca. 450 Ma) William Lake
1506:
666:
after "a brief courtship". Only males care for laid eggs and young.
596:
In total, pycnogonids have four to six pairs of legs for walking. A
3445:
3213:
3200:
1298:, the oldest known pycnogonid, from the Cambrian 'Orsten' of Sweden
3567:
2727:
1287:
750:
710:
668:
655:
638:
543:
462:
408:
1981:
Five
Kingdoms, An Illustrated Guide to the Phyla of Life on Earth
439:
would place them closer to true spiders than to other well-known
3432:
2132:
Brenneis, Georg; Ungerer, Petra; Scholtz, Gerhard (2008-10-27).
1395:
835:
831:
696:
590:
3068:
2767:
521:) with six leg pairs. Pycnogonids do not require a traditional
725:
These animals live in many different parts of the world, from
552:
Certain pycnogonids are so small that each of their very tiny
3187:
2648:
deposit in Manitoba, Canada, is also the first reported from
1582:, Australian Antarctic Division. 22 July 2010. Archived from
2973:
Malacostraca (woodlice, shrimps, crayfish, lobsters, crabs)
2539:"WoRMS - World Register of Marine Species - Stiripasterida"
2181:
Journal of Zoological Systematics and Evolutionary Research
1737:. Dordrecht, NL: Springer Netherlands. pp. 3321–3335.
382:
376:
2515:"WoRMS - World Register of Marine Species - Eupantopodida"
370:
2758:
373:
358:
2998:
Branchiopoda (fairy, tadpole, clam shrimps, water fleas)
2320:
Proceedings of the Royal Society B: Biological Sciences
1576:"Sea spiders provide insights into Antarctic evolution"
814:
Sea spiders have long been considered to belong to the
2563:"WoRMS - World Register of Marine Species - Pantopoda
2491:"WoRMS - World Register of Marine Species - Pantopoda"
1369:
In 2013, the first fossil pycnogonid found within an
509:) with five leg pairs and two species in two genera (
388:
379:
367:
361:
2754:
Images of Pycnogonida, and Pycnogonids in literature
385:
355:
3335:
3077:
3011:
2941:
2932:
2899:
2890:
2863:
2841:
811:are considered part of the single order Pantopoda.
477:includes species with five pairs, and the families
364:
1978:
913:, the order Pantopoda is subdivided as follows:
1154:, including the following genera and families:
901:is the sister group to the remaining families.
533:allows them to suck nutrients from soft-bodied
1398:in south-eastern France. Researchers from the
1161:Sabroux, Edgecombe, Pisani & Garwood, 2023
2779:
8:
3629:Taxa named by Carl Eduard Adolph Gerstaecker
2878:Arachnida (spiders, scorpions, ticks, mites)
2641:
1985:(third ed.). W.H. Freeman and Company.
1784:"Respiratory gut peristalsis by sea spiders"
1777:
1775:
1773:
1407:
783:. Although they can feed by inserting their
1910:Invertebrate Reproduction & Development
1554:
1138:Fry, 1978, including the following family:
3065:
2938:
2896:
2847:
2786:
2772:
2764:
1845:
1843:
873:oceans. However, a subsequent study using
111:
31:
2466:
2405:
2347:
2282:
2233:
2192:
1884:
1807:
1674:
1461:: "taxonomic synonym of Pycnogonida <
921:, including the following superfamilies:
556:consists of a single cell, surrounded by
1580:Department of the Environment and Energy
1422:
593:and tubercle, which projects dorsally.
2759:Bibliography (compiled by Franz Krapp)
580:, and the first pair of walking legs.
435:, their traditional classification as
2978:Thecostraca (barnacles and relatives)
2427:
2425:
1179:Bergstrom, Sturmer & Winter, 1980
7:
3573:6A4A87C4-FFDE-FFC9-D3F6-FC45FDECFC6C
3534:d175446b-2aea-4634-9ee9-fe3227f9d402
3276:faa5c016-9c02-4d6f-a262-4d50baaac9e9
795:rather than a predator of anemones.
427:Although "sea spiders" are not true
2665:"Fossil sea spiders thrill experts"
2372:(2022-02-03). Teeling, Emma (ed.).
846:, among other lesser-known orders.
293:
3031:Diplura (two-pronged bristletails)
1951:McClain, Craig (August 14, 2006).
25:
3624:Extant Cambrian first appearances
2993:Cephalocarida (horseshoe shrimps)
753:that are found along shorelines.
335:
2714:
2702:
2687:
2203:10.1111/j.1439-0469.2004.00284.x
2150:10.1111/j.1525-142X.2008.00285.x
1436:World Register of Marine Species
980:Kim & Hong, 1987 (uncertain)
911:World Register of Marine Species
658:, while males possess a pair of
548:Anatomy of a pantopod pycnogonid
537:, and their digestive tract has
455:to all other living arthropods.
351:
177:
2749:Introduction to the Pycnogonida
2439:Molecular Biology and Evolution
2378:Molecular Biology and Evolution
2222:Molecular Biology and Evolution
1663:Molecular Biology and Evolution
600:and a much smaller unsegmented
1743:10.1007/978-1-4020-6359-6_4098
1553:Pycnogonida, class name, from
1526:Merriam-Webster.com Dictionary
1493:Merriam-Webster.com Dictionary
1456:Merriam-Webster.com Dictionary
338:‘all feet’), belonging to the
120:Left to right, top to bottom:
1:
1930:10.1080/07924259.2003.9652540
564:. The first of these are the
345:, hence they are also called
1977:; Schwartz, Karlene (1998).
1295:Cambropycnogon klausmuelleri
1292:Reconstruction of the larva
1170:Poschmann & Dunlop, 2005
654:. Females possess a pair of
635:Reproduction and development
2963:Pentastomida (tongue worms)
2873:Xiphosura (horseshoe crabs)
2138:Evolution & Development
1698:Ruppert, Edward E. (1994).
858:, the anterior part of the
3645:
1735:Encyclopedia of Entomology
988:Nakamura & Child, 1983
27:Order of marine arthropods
3052:
2855:Pycnogonida (sea spiders)
2850:
2803:
2744:World list of Pycnogonida
1877:10.1016/j.cub.2017.01.047
1809:10.1016/j.cub.2017.05.062
541:extending into the legs.
468:Callipallene brevirostris
308:
301:
290:
285:
267:
260:
174:Scientific classification
172:
119:
110:
34:
3026:Collembola (springtails)
2953:Ostracoda (seed shrimps)
2638:Palaeomarachne granulata
707:Distribution and ecology
3057:are paraphyletic groups
2735:Encyclopædia Britannica
2596:Journal of Paleontology
2433:Crandall, Keith (ed.).
1321:Coalbrookdale Formation
644:Tanystylum californicum
629:open circulatory system
2968:Branchiura (fish lice)
2924:Diplopoda (millipedes)
2909:Chilopoda (centipedes)
2738:(11th ed.). 1911.
2642:
2451:10.1093/molbev/msaa228
2390:10.1093/molbev/msac021
2332:10.1098/rspb.2019.2426
1555:
1408:
1299:
722:
677:
664:external fertilisation
647:
549:
470:
3555:Paleobiology Database
3297:Paleobiology Database
2644:Konservat-Lagerstätte
2284:10.1093/sysbio/syz011
2235:10.1093/molbev/msu235
1676:10.1093/molbev/msu235
1291:
885:(consistent with the
714:
672:
642:
547:
466:
41:Late Cambrian–present
2798:classes by subphylum
2723:at Wikimedia Commons
1701:Invertebrate Zoology
1379:) was reported from
1159:Palaeopycnogonididae
625:peristaltic movement
146:Ammothea hilgendorfi
3021:Protura (coneheads)
2729:"Pycnogonida"
2608:2013JPal...87..395R
2079:10.1038/nature04591
2071:2006Natur.441..506J
2028:10.1038/nature03984
2020:2005Natur.437.1144M
1922:2003InvRD..43..193B
1869:2017CBio...27..638B
1800:2017CBio...27.R638W
1628:10.1038/nature08742
1620:2010Natur.463.1079R
1545:The Free Dictionary
1498:: "New Latin, from
1392:La Voulte-sur-Rhône
716:Nymphon leptocheles
138:Pycnogonum stearnsi
122:Austrodecus bamberi
2673:. August 16, 2007.
2326:(1917): 20192426.
2271:Systematic Biology
1957:Deep Sea News Info
1529:. Merriam-Webster.
1496:. Merriam-Webster.
1459:. Merriam-Webster.
1400:University of Lyon
1300:
723:
678:
648:
568:, followed by the
550:
523:respiratory system
471:
406:; with the suffix
3606:
3605:
3542:Open Tree of Life
3284:Open Tree of Life
3071:Taxon identifiers
3062:
3061:
3048:
3047:
3044:
3043:
3036:Insecta (insects)
2886:
2885:
2719:Media related to
2695:Arthropods portal
2571:marinespecies.org
2543:marinespecies.org
2519:marinespecies.org
2495:marinespecies.org
2228:(11): 2963–2984.
1992:978-0-7167-3027-9
1794:(13): R638–R639.
1752:978-1-4020-6359-6
1614:(7284): 1079–83.
1578:(Press release).
1111:Phoxichilidioidea
1081:Rhynchothoracidae
909:According to the
826:, which includes
743:Mediterranean Sea
558:connective tissue
494:Pentacolossendeis
318:
317:
313:
256:
239:
16:(Redirected from
3636:
3599:
3598:
3586:
3585:
3576:
3575:
3563:
3562:
3550:
3549:
3537:
3536:
3527:
3526:
3514:
3513:
3511:NHMSYS0021049490
3501:
3500:
3488:
3487:
3475:
3474:
3462:
3461:
3449:
3448:
3436:
3435:
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3314:
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3292:
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3279:
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3216:
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3112:
3111:
3098:
3097:
3096:
3066:
2939:
2897:
2848:
2788:
2781:
2774:
2765:
2739:
2731:
2718:
2707:Data related to
2706:
2697:
2692:
2691:
2675:
2674:
2661:
2655:
2654:
2647:
2632:
2630:
2616:10.1666/12-057.1
2587:
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2580:
2578:
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2559:
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2311:
2305:
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2286:
2262:
2256:
2255:
2237:
2213:
2207:
2206:
2196:
2176:
2170:
2169:
2129:
2123:
2122:
2120:
2119:
2113:scienceblogs.com
2105:
2099:
2098:
2054:
2048:
2047:
2014:(7062): 1144–8.
2003:
1997:
1996:
1984:
1971:
1965:
1964:
1959:. Archived from
1948:
1942:
1941:
1905:
1899:
1898:
1888:
1847:
1838:
1837:
1811:
1779:
1768:
1763:
1757:
1756:
1730:
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1723:
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1688:
1678:
1654:
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1572:
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1558:
1537:
1531:
1530:
1517:
1511:
1497:
1484:
1478:
1460:
1447:
1441:
1440:
1439:. Taxon details.
1427:
1413:
1339:Flagellopantopus
1237:Philippi, 1842 (
1167:Flagellopantopus
1125:Phoxichilidiidae
818:, together with
443:groups, such as
395:
394:
391:
390:
387:
384:
381:
378:
375:
372:
369:
366:
363:
360:
357:
337:
311:
251:
234:
182:
181:
162:Nymphon signatum
115:
105:
42:
38:Temporal range:
32:
21:
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3643:
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3517:
3509:
3504:
3496:
3491:
3483:
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3470:
3465:
3457:
3452:
3444:
3439:
3431:
3426:
3418:
3413:
3405:
3400:
3392:
3387:
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3365:
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3147:
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3086:
3073:
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3040:
3007:
2928:
2882:
2859:
2837:
2799:
2792:
2726:
2693:
2686:
2683:
2678:
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2658:
2628:
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2589:
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2513:
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2431:
2430:
2423:
2370:
2369:
2365:
2313:
2312:
2308:
2264:
2263:
2259:
2215:
2214:
2210:
2194:10.1.1.714.8297
2178:
2177:
2173:
2131:
2130:
2126:
2117:
2115:
2107:
2106:
2102:
2065:(7092): 506–8.
2056:
2055:
2051:
2005:
2004:
2000:
1993:
1973:
1972:
1968:
1963:on 9 July 2007.
1950:
1949:
1945:
1907:
1906:
1902:
1857:Current Biology
1849:
1848:
1841:
1788:Current Biology
1781:
1780:
1771:
1764:
1760:
1753:
1732:
1731:
1727:
1712:
1697:
1696:
1692:
1669:(11): 2963–84.
1656:
1655:
1651:
1604:
1603:
1599:
1589:
1587:
1586:on 31 July 2018
1574:
1573:
1569:
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1538:
1534:
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1514:
1486:
1485:
1481:
1449:
1448:
1444:
1429:
1428:
1424:
1420:
1286:
1249:Goodsir, 1842 (
1096:Callipallenidae
1069:Jarzynsky, 1870
1060:Colossendeoidea
947:Ascorhynchoidea
907:
860:arthropod brain
820:horseshoe crabs
801:
709:
675:Achelia spinosa
637:
461:
354:
350:
281:
275:
250:
233:
176:
154:Endeis flaccida
106:
104:
103:
98:
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88:
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78:
73:
68:
63:
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53:
48:
40:
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22:
15:
12:
11:
5:
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2793:
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2783:
2776:
2768:
2762:
2761:
2756:
2751:
2746:
2740:
2724:
2712:
2711:at Wikispecies
2699:
2698:
2682:
2681:External links
2679:
2677:
2676:
2656:
2602:(3): 395–405.
2582:
2565:incertae sedis
2554:
2530:
2506:
2482:
2445:(2): 686–701.
2421:
2384:(2): msac021.
2363:
2306:
2277:(6): 896–917.
2257:
2208:
2171:
2144:(6): 717–724.
2124:
2100:
2049:
1998:
1991:
1975:Margulis, Lynn
1966:
1943:
1916:(3): 193–222.
1900:
1863:(5): 638–650.
1839:
1769:
1758:
1751:
1725:
1710:
1690:
1649:
1597:
1567:
1532:
1512:
1479:
1442:
1421:
1419:
1416:
1376:Palaeomarachne
1345:Palaeopantopus
1334:Hunsrück Slate
1313:Cambropycnogon
1285:
1282:
1281:
1280:
1279:
1278:
1266:
1254:
1242:
1230:
1218:
1206:
1194:
1180:
1171:
1162:
1152:incertae sedis
1148:
1147:
1146:
1136:Stiripasterida
1132:
1131:
1130:
1129:
1128:
1121:
1108:
1107:
1106:
1099:
1086:
1085:
1084:
1083:Thompson, 1909
1077:
1070:
1067:Colossendeidae
1057:
1056:
1055:
1054:
1053:
1045:
1037:
1029:
1021:
1013:
1005:
1004:Hedgpeth, 1943
997:
989:
981:
973:
962:incertae sedis
957:
954:Ascorhynchidae
944:
943:
942:
935:
906:
905:Group taxonomy
903:
800:
799:Classification
797:
708:
705:
652:hermaphroditic
636:
633:
479:Colossendeidae
460:
457:
396:; named after
316:
315:
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3258:
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3171:
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3154:
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3141:
3137:
3132:
3128:
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3119:
3115:
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3104:
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3019:
3018:
3016:
3014:
3010:
3004:
3001:
2999:
2996:
2994:
2991:
2989:
2988:Tantulocarida
2986:
2984:
2981:
2979:
2976:
2974:
2971:
2969:
2966:
2964:
2961:
2959:
2958:Mystacocarida
2956:
2954:
2951:
2950:
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2940:
2937:
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2925:
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2898:
2895:
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2868:
2866:
2865:Euchelicerata
2862:
2856:
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2840:
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2828:
2824:
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2159:
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2128:
2125:
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2110:
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2076:
2072:
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2053:
2050:
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2041:
2037:
2033:
2029:
2025:
2021:
2017:
2013:
2009:
2002:
1999:
1994:
1988:
1983:
1982:
1976:
1970:
1967:
1962:
1958:
1954:
1953:"Sea Spiders"
1947:
1944:
1939:
1935:
1931:
1927:
1923:
1919:
1915:
1911:
1904:
1901:
1896:
1892:
1887:
1882:
1878:
1874:
1870:
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1801:
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1778:
1776:
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1759:
1754:
1748:
1744:
1740:
1736:
1729:
1726:
1721:
1717:
1713:
1711:0-03-026668-8
1707:
1703:
1702:
1694:
1691:
1686:
1682:
1677:
1672:
1668:
1664:
1660:
1653:
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1527:
1522:
1516:
1513:
1509:
1508:
1503:
1502:
1495:
1494:
1489:
1488:"Pycnogonida"
1483:
1480:
1476:
1475:
1470:
1469:
1464:
1458:
1457:
1452:
1446:
1443:
1438:
1437:
1432:
1431:"Pycnogonida"
1426:
1423:
1417:
1415:
1412:
1411:
1405:
1401:
1397:
1393:
1388:
1386:
1382:
1378:
1377:
1372:
1367:
1365:
1364:
1363:Pentapantopus
1359:
1358:
1353:
1352:
1347:
1346:
1341:
1340:
1335:
1332:
1328:
1327:
1322:
1319:
1315:
1314:
1310:' of Sweden (
1309:
1305:
1297:
1296:
1290:
1284:Fossil record
1283:
1276:
1273:Costa, 1861 (
1272:
1271:
1267:
1264:
1261:Costa, 1836 (
1260:
1259:
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1248:
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1236:
1235:
1231:
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1225:Hesse, 1874 (
1224:
1223:
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1211:
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1201:Costa, 1836 (
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1160:
1156:
1155:
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1144:
1143:Austrodecidae
1140:
1139:
1137:
1133:
1126:
1122:
1119:
1115:
1114:
1112:
1109:
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1092:
1091:Pocock, 1904
1090:
1087:
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1063:
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1058:
1052:Barnard, 1946
1051:
1050:
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1035:
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1025:Pigrogromitus
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949:Pocock, 1904
948:
945:
940:
939:Pallenopsidae
936:
933:
929:
928:
926:
923:
922:
920:
919:Eupantopodida
916:
915:
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904:
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899:Austrodecidae
896:
892:
888:
884:
879:
876:
872:
868:
867:
861:
857:
856:protocerebrum
853:
852:deutocerebrum
847:
845:
841:
837:
833:
829:
825:
821:
817:
812:
810:
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786:
782:
778:
774:
770:
767:that feed on
766:
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748:
747:Caribbean Sea
744:
740:
739:United States
737:coast of the
736:
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718:grazing on a
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583:
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425:
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412:). They are
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3054:
2942:
2934:Pancrustacea
2854:
2831:Superphylum
2733:
2668:
2659:
2637:
2634:
2629:23 September
2627:. Retrieved
2599:
2595:
2585:
2574:. Retrieved
2570:
2564:
2557:
2546:. Retrieved
2542:
2533:
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2494:
2485:
2442:
2438:
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2180:
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2127:
2116:. Retrieved
2112:
2103:
2062:
2058:
2052:
2011:
2007:
2001:
1980:
1969:
1961:the original
1956:
1946:
1913:
1909:
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1734:
1728:
1700:
1693:
1666:
1662:
1652:
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1607:
1600:
1588:. Retrieved
1584:the original
1570:
1548:
1544:
1541:"pycnogonid"
1535:
1524:
1521:"pycnogonid"
1515:
1505:
1499:
1491:
1482:
1472:
1466:
1454:
1445:
1434:
1425:
1389:
1381:William Lake
1374:
1368:
1361:
1355:
1351:Palaeoisopus
1349:
1343:
1337:
1336:of Germany (
1324:
1323:of England (
1311:
1301:
1293:
1275:nomen dubium
1274:
1268:
1263:nomen dubium
1262:
1256:
1251:nomen dubium
1250:
1244:
1239:nomen dubium
1238:
1232:
1227:nomen dubium
1226:
1220:
1215:nomen dubium
1214:
1208:
1203:nomen dubium
1202:
1196:
1190:nomen dubium
1188:
1182:
1174:
1165:
1151:
1120:Norman, 1908
1105:Wilson, 1878
1098:Hilton, 1942
1089:Nymphonoidea
1076:Wilson, 1878
1074:Pycnogonidae
1047:
1039:
1033:Pycnopallene
1031:
1028:Calman, 1927
1023:
1015:
1007:
999:
996:Hilton, 1939
991:
983:
977:Bradypallene
975:
972:Bamber, 2004
967:
960:
927:Dohrn, 1881
925:Ammotheoidea
908:
880:
866:Anomalocaris
864:
848:
813:
807:exists. All
802:
789:sea anemones
755:
724:
715:
690:
686:
679:
674:
649:
643:
614:
595:
551:
516:
512:Dodecolopoda
510:
506:Pentanymphon
504:
498:
492:
486:
475:Pycnogonidae
472:
467:
453:sister group
437:chelicerates
426:
414:cosmopolitan
407:
397:
346:
342:
331:
320:
319:
310:Arachnopoda
309:
291:
269:
268:
247:
230:
161:
153:
145:
142:Pycnogonidae
137:
130:Colossendeis
129:
121:
29:
3619:Pycnogonids
3467:iNaturalist
3361:Wikispecies
3222:iNaturalist
3136:Pycnogonida
3123:Pycnogonida
3109:Pycnogonida
3103:Wikispecies
3079:Pycnogonida
2892:Mandibulata
2843:Chelicerata
2827:Protostomia
2825:(unranked)
2819:(unranked)
2813:Subkingdom
2742:PycnoBase:
2721:Pycnogonida
2709:Pycnogonida
1590:27 December
1451:"Pantopoda"
1410:Lagerstätte
1394:, south of
1270:Platychelus
1210:Oiceobathys
1145:Stock, 1954
1113:Sars, 1891
1103:Nymphonidae
1044:Loman, 1921
1036:Stock, 1950
1020:Child, 1982
1017:Mimipallene
1001:Ephyrogymna
934:Dohrn, 1881
932:Ammotheidae
883:Chelicerata
816:Chelicerata
777:polychaetes
758:carnivorous
731:New Zealand
693:polychaetes
610:chelicerate
606:exoskeleton
582:Nymphonidae
539:diverticula
518:Sexanymphon
500:Pentapycnon
483:Nymphonidae
459:Description
449:crustaceans
431:, nor even
422:deep waters
347:pycnogonids
343:Pycnogonida
323:are marine
321:Sea spiders
253:Gerstaecker
231:Pycnogonida
221:Chelicerata
217:Subphylum:
166:Nymphonidae
150:Ammotheidae
35:Sea spiders
18:Sea spiders
3613:Categories
2796:Arthropoda
2576:2023-12-04
2548:2023-12-04
2524:2023-12-04
2500:2023-12-04
2118:2022-01-10
1561:type genus
1556:Pycnogonum
1501:Pycnogonum
1418:References
1406:La Voulte
1371:Ordovician
1357:Palaeothea
1329:) and the
1258:Phanodemus
1198:Foxichilus
1176:Palaeothea
1127:Sars, 1891
1012:Hoek, 1881
956:Hoek, 1881
844:harvestmen
822:, and the
769:cnidarians
765:scavengers
733:, and the
566:chelifores
562:appendages
488:Decolopoda
404:type genus
399:Pycnogonum
325:arthropods
312:Dana, 1853
271:Pycnogonum
262:Type genus
211:Arthropoda
3381:Pantopoda
3367:Pantopoda
3337:Pantopoda
3003:Remipedia
2944:Crustacea
2919:Pauropoda
2901:Myriapoda
2833:Ecdysozoa
2821:Bilateria
2815:Eumetazoa
2650:Laurentia
2459:1537-1719
2398:0737-4038
2340:0962-8452
2293:1063-5157
2244:1537-1719
2189:CiteSeerX
1818:0960-9822
1551:Neo-Latin
1463:Neo-Latin
1373:deposit (
1326:Haliestes
1246:Pephredro
1150:suborder
1134:suborder
1041:Pycnothea
993:Decachela
985:Chonothea
941:Fry, 1978
917:suborder
893:putative
891:chelicera
887:chelifore
840:scorpions
824:Arachnida
785:proboscis
781:bryozoans
761:predators
741:, to the
727:Australia
701:ovigerous
673:Larva of
646:with eggs
617:hemolymph
587:proboscis
578:courtship
531:proboscis
527:diffusion
441:arthropod
433:arachnids
418:Antarctic
332:Pantopoda
286:Families
248:Pantopoda
236:Latreille
197:Kingdom:
191:Eukaryota
3402:BugGuide
3346:Wikidata
3157:BugGuide
3088:Wikidata
3013:Hexapoda
2983:Copepoda
2914:Symphyla
2809:Animalia
2807:Kingdom
2670:BBC News
2624:83924778
2477:32915961
2416:35137183
2358:31847768
2301:30917194
2252:25107551
2187:: 8–21.
2158:19021742
2087:16724066
2036:16237442
1938:84345599
1895:28238655
1834:35014992
1826:28697358
1720:30544625
1685:25107551
1636:20147900
1404:Jurassic
1385:Manitoba
1331:Devonian
1318:Silurian
1304:Cambrian
1234:Paritoca
1184:Alcynous
1118:Endeidae
1009:Hannonia
895:homology
875:Hox gene
871:Cambrian
809:families
793:parasite
745:and the
303:Synonyms
278:Brünnich
207:Phylum:
201:Animalia
187:Domain:
158:Endeinae
3459:9477065
3433:2598723
3352:Q874792
2794:Extant
2604:Bibcode
2468:7826184
2407:8845124
2349:6939912
2166:6048195
2095:4307398
2067:Bibcode
2044:4400419
2016:Bibcode
1918:Bibcode
1886:5344685
1865:Bibcode
1796:Bibcode
1644:4427443
1616:Bibcode
1465:, from
1222:Oomerus
1141:Family
1123:Family
1116:Family
1101:Family
1094:Family
1079:Family
1072:Family
1065:Family
1049:Queubus
959:Family
952:Family
937:Family
930:Family
828:spiders
773:sponges
735:Pacific
720:hydroid
682:ovigers
656:ovaries
602:abdomen
574:ovigers
554:muscles
445:insects
429:spiders
327:of the
244:Order:
227:Class:
3583:516419
3580:uBio:
3547:247585
3531:NZOR:
3524:373319
3472:245099
3446:1PANTO
3407:810330
3312:516418
3309:uBio:
3289:910210
3273:NZOR:
3201:1PYCNC
3162:810329
3094:Q19106
3055:italic
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2059:Nature
2042:
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2008:Nature
1989:
1936:
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1883:
1832:
1824:
1816:
1749:
1718:
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1683:
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1634:
1608:Nature
1308:Orsten
842:, and
805:orders
779:, and
660:testes
572:, the
503:, and
402:, the
280:, 1764
255:, 1863
238:, 1810
3591:WoRMS
3568:Plazi
3560:19020
3498:83546
3485:10867
3480:IRMNG
3420:8ZXHL
3394:80508
3320:WoRMS
3302:19015
3266:57294
3253:83545
3235:IRMNG
3227:47825
3149:26059
2620:S2CID
2162:S2CID
2091:S2CID
2040:S2CID
1934:S2CID
1830:S2CID
1640:S2CID
1549:From
1474:-poda
1468:pant-
969:Bango
836:ticks
832:mites
787:into
751:algae
697:clams
621:moult
570:palps
340:class
329:order
132:sp. (
3596:1358
3519:NCBI
3493:ITIS
3454:GBIF
3441:EPPO
3389:BOLD
3325:1302
3261:NCBI
3248:ITIS
3240:1325
3209:GBIF
3196:EPPO
3188:1547
3144:BOLD
2631:2017
2473:PMID
2455:ISSN
2412:PMID
2394:ISSN
2354:PMID
2336:ISSN
2297:PMID
2289:ISSN
2248:PMID
2240:ISSN
2154:PMID
2083:PMID
2032:PMID
1987:ISBN
1891:PMID
1822:PMID
1814:ISSN
1747:ISBN
1716:OCLC
1706:ISBN
1681:PMID
1632:PMID
1592:2017
1507:-ida
1396:Lyon
1383:in
1360:and
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591:anus
585:the
529:. A
515:and
481:and
420:and
336:lit.
294:text
292:See
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2402:PMC
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2328:doi
2324:286
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2199:doi
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