Sea spider

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Sea spiders
Temporal range: Late Cambrian–present
Sea spider.jpg
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Chelicerata
Class: Pycnogonida
Latreille, 1810
Order: Pantopoda
Gerstaecker, 1863

Sea spiders are marine arthropods of the order Pantopoda [1] (lit. ‘all feet’ [2] ), belonging to the class Pycnogonida, [3] hence they are also called pycnogonids (named after Pycnogonum , the type genus; [4] with the suffix -id ). They are cosmopolitan, found in oceans around the world. There are over 1,300 known species, with legs ranging from 1 mm (0.04 in) to over 70 cm (2.3 ft). [5] Most are toward the smaller end of this range in relatively shallow depths; however, they can grow to be quite large in Antarctic and deep waters.

Contents

Although "sea spiders" are not true spiders, or even arachnids, their traditional classification as chelicerates would place them closer to true spiders than to other well-known arthropod groups, such as insects or crustaceans. This is in dispute, however, as genetic evidence suggests they may be the sister group to all other living arthropods. [6] [7]

Description

Anatomy of a pycnogonid: A: head; B: thorax; C: abdomen 1: proboscis; 2: chelifores; 3: palps; 4: ovigers; 5: egg sacs; 6a-6d: four pairs of legs Pycnogonida Nymphon s Sars.png
Anatomy of a pycnogonid: A: head; B: thorax; C: abdomen 1: proboscis; 2: chelifores; 3: palps; 4:  ovigers; 5: egg sacs; 6a–6d: four pairs of legs

Sea spiders have long legs in contrast to a small body size. The number of walking legs is usually eight (four pairs), but species with five and six pairs exist. Pycnogonids do not require a traditional respiratory system. Instead, gasses are absorbed by the legs and transferred through the body by diffusion. A proboscis allows them to suck nutrients from soft-bodied invertebrates, and their digestive tract has diverticula extending into the legs.

A pycnogonid grazing on a hydroid Nymphon-leptocheles.jpg
A pycnogonid grazing on a hydroid

Certain pycnogonids are so small that each of their very tiny muscles consists of only one single cell, surrounded by connective tissue. The anterior region consists of the proboscis, which has fairly limited dorsoventral and lateral movement, and three to four appendages including the ovigers, which are used in caring for young and cleaning as well as courtship. In some species, the chelifores, palps and ovigers can be reduced or missing in adults. In those species that lack chelifores and palps, the proboscis is well developed and more mobile and flexible, often equipped with numerous sensory bristles and strong rasping ridges around the mouth. The last segment includes the anus and tubercle, which projects dorsally.

In total, pycnogonids have four to six pairs of legs for walking as well as other appendages which often resemble legs. A cephalothorax and much smaller abdomen make up the extremely reduced body of the pycnogonid, which has up to two pairs of dorsally located simple eyes on its non-calcareous exoskeleton, 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 chelicerate extend throughout many appendages because its body is too small to accommodate all of them alone.

The morphology of the sea spider creates an extremely well suited surface-area to volume ratio for any respiration to occur through direct diffusion. Oxygen is absorbed by the legs and is transported via the hemolymph to the rest of the body. [8] The most recent research seems to indicate that waste leaves the body through the digestive tract or is lost during a moult. 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. [8] Hemolymph circulation in the legs is mostly driven by the peristaltic movement in the part of the gut that extends into every leg. [8] These creatures possess an open circulatory system 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.

Reproduction and development

All pycnogonid species have separate sexes, except for one species that is hermaphroditic. Females possess a pair of ovaries, while males possess a pair of testes located dorsally in relation to the digestive tract. Reproduction involves external fertilisation after "a brief courtship". Only males care for laid eggs and young.

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 ovigers. 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.

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 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.

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 polychaetes and clams. When the attaching larva hatches it still looks like an embryo, and immediately attaches itself to the ovigerous 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.

Distribution and ecology

A pycnogonid in its natural habitat Pycnogonid.jpg
A pycnogonid in its natural habitat

These animals live in many different parts of the world, from Australia, New Zealand, and the Pacific coast of the United States, to the Mediterranean Sea and the Caribbean Sea, 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 algae that are found along shorelines.

Sea spiders either walk along the bottom with their stilt-like legs or swim just above it using an umbrella pulsing motion. [9] Sea spiders are mostly carnivorous predators or scavengers that feed on cnidarians, sponges, polychaetes, and bryozoans. Although they can feed by inserting their proboscis into sea anemones, which are much larger, most sea anemones survive this ordeal, making the sea spider a parasite rather than a predator of anemones.

Classification

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 orders exists. All families are considered part of the single order Pantopoda.

Sea spiders have long been considered to belong to the Chelicerata, together with horseshoe crabs, and the Arachnida, which includes spiders, mites, ticks, scorpions, and harvestmen, among other, lesser-known orders. [10]

A competing theory proposes that pycnogonida belong to their own lineage, distinct from chelicerates, crustaceans, myriapods, or insects. This theory contends that the sea spider's chelifores, which are unique among extant arthropods, are not in any way homologous to the chelicerae in real chelicerates, as was previously supposed. Instead of developing from the deutocerebrum, they can be traced to the protocerebrum, the anterior part of the arthropod brain and found in the first head segment that in all other arthropods give rise to the eyes only. This is not found anywhere else among arthropods, except in some fossil forms like Anomalocaris , indicating that the 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 is confirmed, it would mean the sea spiders are the last surviving (and highly modified) members of an ancient stem group of arthropods that lived in Cambrian oceans. [11] However, a subsequent study using Hox gene expression patterns consistent with a developmental homology between chelicerates and chelifores, with chelifores innervated from a deuterocerebrum that has been rotated forwards; thus, the protocerebral Great Appendage clade does not include the Pycnogonida. [12] [13]

Recent work places the Pycnogonida outside the Arachnomorpha as basal Euarthropoda, or inside Chelicerata (based on the chelifore-chelicera putative homology). [14]

Group taxonomy

According to the World Register of Marine Species, the order Pantopoda is subdivided as follows: [15]

This taxonomic classification replaces the older version in which Pantopoda is subdivided into families as follows:

Fossil record

Although the fossil record of pycnogonids is scant, it is clear that they once possessed a coelom, but it was later lost, and that the group is very old.[ citation needed ]

The earliest fossils are known from the Cambrian 'Orsten' of Sweden, the Silurian Wenlock Series of England and the Devonian Hunsrück Slate of Germany. 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.

In 2013, the first fossil pycnogonid found within an Ordovician deposit was reported from William Lake in northern Manitoba. [19]

In 2007, remarkably well preserved fossils were exposed in fossil beds at La Voulte-sur-Rhône, south of Lyon in south-eastern France. Researchers from the University of Lyon discovered about 70 fossils from three distinct species in the 160-million-year-old Jurassic La Voulte Lagerstätte . The find will help fill in an enormous gap in the history of these creatures. [20]

Related Research Articles

Chelicerata Subphylum of arthropods

The subphylum Chelicerata constitutes one of the major subdivisions of the phylum Arthropoda. It contains the sea spiders, arachnids, and several extinct lineages, such as the eurypterids and chasmataspidids.

Arachnid Class of arthropods

Arachnida is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Arachnida includes orders containing spiders, scorpions, ticks, mites, harvestmen, and solifuges. In 2019, a molecular phylogenetic study also placed horseshoe crabs in Arachnida.

Eurypterid Order of arthropods (fossil)

Eurypterids, often informally called sea scorpions, are a group of extinct arthropods that form the order Eurypterida. The earliest known eurypterids date to the Darriwilian stage of the Ordovician period 467.3 million years ago. The group is likely to have appeared first either during the Early Ordovician or Late Cambrian period. With approximately 250 species, the Eurypterida is the most diverse Paleozoic chelicerate order. Following their appearance during the Ordovician, eurypterids became major components of marine faunas during the Silurian, from which the majority of eurypterid species have been described. The Silurian genus Eurypterus accounts for more than 90% of all known eurypterid specimens. Though the group continued to diversify during the subsequent Devonian period, the eurypterids were heavily affected by the Late Devonian extinction event. They declined in numbers and diversity until becoming extinct during the Permian–Triassic extinction event 251.9 million years ago.

Xiphosura Order of marine chelicerates

Xiphosura is an order of arthropods related to arachnids. They are sometimes called horseshoe crabs. They first appeared in the Hirnantian. Currently, there are only four living species. Xiphosura contains one suborder, Xiphosurida, and several stem-genera.

Arachnomorpha

Arachnomorpha is a subdivision or clade of Arthropoda, comprising the monophyletic group formed by the trilobites, other great appendage arthropods and trilobite-like families, and a diverse sister clade including the chelicerates. Great debate is held on the position of the Pycnogonida, which are currently thought not to be placed in the immediate vicinity of the Chelicerata. Arachnomorpha are considered the sister group to the crustaceans, which are increasingly being accepted as members of the mandibulate clade.

Cheloniellida Order of arthropods (fossil)

Cheloniellida is a taxon of extinct Paleozoic arthropods. As of 2018, 8 monotypic genera of cheloniellids had been formally described, whose fossils are found in marine strata ranging from Ordovician to Devonian in age. Cheloniellida has a controversial phylogenetic position, with previous studies associated it as either a member or relative of various fossil and extant arthropod taxa. It was later accepted as a member of Vicissicaudata within Artiopoda.

Arthropod mouthparts

The mouthparts of arthropods have evolved into a number of forms, each adapted to a different style or mode of feeding. Most mouthparts represent modified, paired appendages, which in ancestral forms would have appeared more like legs than mouthparts. In general, arthropods have mouthparts for cutting, chewing, piercing, sucking, shredding, siphoning, and filtering. This article outlines the basic elements of four arthropod groups: insects, myriapods, crustaceans and chelicerates. Insects are used as the model, with the novel mouthparts of the other groups introduced in turn. Insects are not, however, the ancestral form of the other arthropods discussed here.

Arthropod head problem Uncertainty regarding the evolutionary relationship of the segmental composition of the head in various arthropod groups

The (pan)arthropod head problem is a long-standing zoological dispute concerning the segmental composition of the heads of the various arthropod groups, and how they are evolutionarily related to each other. While the dispute has historically centered on the exact make-up of the insect head, it has been widened to include other living arthropods such as the crustaceans and chelicerates; and fossil forms, such as the many arthropods known from exceptionally preserved Cambrian faunas. While the topic has classically been based on insect embryology, in recent years a great deal of developmental molecular data has become available. Dozens of more or less distinct solutions to the problem, dating back to at least 1897, have been published, including several in the 2000s.

<i>Mixopterus</i>

Mixopterus is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils of Mixopterus have been discovered in deposits from Late Silurian age, and have been referred to several different species. Fossils have been recovered from two continents; Europe and North America.

Chasmataspidid

Chasmataspidids, sometime referred to as chasmataspids, are a group of extinct chelicerate arthropods that form the order Chasmataspidida. Chasmataspidids are probably related to horseshoe crabs (Xiphosura) and/or sea scorpions (Eurypterida), with more recent studies suggest that they form a clade (Dekatriata) with Eurypterida and Arachnida. Chasmataspidids are known sporadically in the fossil record through to the mid-Devonian, with possible evidence suggesting that they were also present during the late Cambrian. Chasmataspidids are most easily recognised by having an opisthosoma divided into a wide forepart (preabdomen) and a narrow hindpart (postabdomen) each comprising 4 and 9 segments respectively. There is some debate about whether they form a natural group.

<i>Maevia inclemens</i> Species of spider

Maevia inclemens is a relatively common and colorful jumping spider of North America. In the males there are two forms, a very rare phenomenon in zoology. These use different courting displays, and differ in appearance: the "tufted" morph has a black body and pedipalps ("palps"), three black tufts across its "head", and pale legs; and the "gray" morph has black and white stripes all over its body and legs, orange palps, and no tufts. However, each form accounts for 50% of the adult males, and they are equally successful in mating. A female of Maevia inclemens is 6.5 to 8.0 millimetres long, while males are 4.75 to 6.50 millimetres long.

Spider Order of arachnids

Spiders are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species diversity among all orders of organisms. Spiders are found worldwide on every continent except for Antarctica, and have become established in nearly every habitat with the exceptions of air and sea colonization. As of July 2019, at least 48,200 spider species, and 120 families have been recorded by taxonomists. However, there has been dissension within the scientific community as to how all these families should be classified, as evidenced by the over 20 different classifications that have been proposed since 1900.

Pycnogonidae Family of sea spiders

Pycnogonidae is a family of sea spiders.

<i>Pycnogonum</i> Genus of sea spiders

Pycnogonum is a genus of sea spiders in the family Pycnogonidae. It is the type genus of the family.

Pycnogonum stearnsi or Stearns' sea spider is a marine arthropod in the family Pycnogonidae. It is found on the western seaboard of North America.

Crustacean Subphylum of arthropods

Crustaceans form a large, diverse arthropod taxon which includes such animals as crabs, lobsters, crayfish, shrimps, prawns, krill, woodlice, and barnacles. The crustacean group can be treated as a subphylum under the clade Mandibulata; because of recent molecular studies it is now well accepted that the crustacean group is paraphyletic, and comprises all animals in the clade Pancrustacea other than hexapods. Some crustaceans are more closely related to insects and the other hexapods than they are to certain other crustaceans.

Nymphon brevirostre is a species of sea spider first described by George Hodge in 1863. The species highly resembles other members of the genus Nymphon, and species identification from morphological traits alone is therefore a complex task.

<i>Nymphon gracile</i> Species of sea spider

Nymphon gracile is a species of sea spider first described by William Elford Leach in 1863. The species highly resembles other members of the genus Nymphon, and species identification from morphological traits alone is, therefore, a complex task.

Palaeoisopus Extinct genus of sea spiders

Palaeoisopus is a monotypic genus of fossil pycnogonid, known only by one species, Palaeoisopus problematicus, discovered from the Lower Devonian Hunsrück Slate of Germany. It have several characters unusual for a pycnogonid, such as swimming legs with alternating size, medially-arranged eyes, and most significantly, a long, segmented abdomen, which were highly reduced in modern counterparts.

<i>Chasmataspis</i>

Chasmataspis is a genus of chasmataspidid, a group of extinct aquatic chelicerate arthropods.

References

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  19. Rudkin, Dave; Cuggy, Michael B.; Young, Graham A.; Thompson, Deborah P. (2013). "An Ordovician Pycnogonid (Sea Spider) with Serially Subdivided 'Head' Region". Journal of Paleontology. 87 (3): 395–405. doi:10.1666/12-057.1. S2CID   83924778 . Retrieved 23 September 2017. 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, Palaeomarachne granulata n. gen. n. sp., from the Upper Ordovician (ca. 450 Ma) William Lake Konservat-Lagerstätte deposit in Manitoba, Canada, is also the first reported from Laurentia. It is the only record thus far of a fossil sea spider in rocks of demonstrably shallow marine origin.
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