Hibbertopteridae

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Hibbertopteridae
Temporal range: Middle Devonian - Late Permian, 387.7–251.9  Ma
A monograph of the British fossil Crustacea (Pl. XXV) (7394041678).jpg
Drawings of the carapace of Hibbertopterus scouleri , from above (upper drawing) and below (lower drawing) and of the ornamentation of the posterior side of the head (enlarged, left) by Henry Woodward, 1866-1878.
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Chelicerata
Order: Eurypterida
Suborder: Stylonurina
Superfamily: Mycteropoidea
Family: Hibbertopteridae
Kjellesvig-Waering, 1959
Type genus
Hibbertopterus
Kjellesvig-Waering, 1959
Genera
Synonyms
  • Cyrtoctenidae
    Waterston et al., 1985

Hibbertopteridae (the name deriving from the type genus Hibbertopterus , meaning "Hibbert's wing") is a family of eurypterids, an extinct group of aquatic arthropods. They were members of the superfamily Mycteropoidea. Hibbertopterids were large, broad and heavy animals unlike virtually every other group of eurypterids, which are commonly streamlined and lightweight. Their bizarre morphology is so unusual that they in the past have been thought to represent an entirely distinct order of chelicerates. Fossils of the family first appear in deposits of Middle Devonian age and the last known fossils representing hibbertopterids are known from deposits of Late Permian age. The hibbertopterids represent the last known living eurypterids, going extinct during the Permian–Triassic extinction event or shortly before. [1]

Contents

Although eurypterids are commonly known as "sea scorpions", hibbertopterids inhabited freshwater swamps and rivers and were unable to swim since they lacked swimming paddles, a feature they shared with their entire suborder, the Stylonurina. Hibbertopterids fed using a method referred to as sweep-feeding, in which the animal would rake through the soft sediment of their substrate with specialised blades on their forward-facing appendages to capture small invertebrates. Though this method of feeding was present in other mycteropoids and in stylonuroids, it was at its most advanced stage within the derived Hibbertopteridae.

The hibbertopterids were the largest of all stylonurine eurypterids, with both Hibbertopterus at 180–200 centimetres (5.9–6.6 ft) and Campylocephalus at 140 centimetres (4.6 feet) representing giant eurypterids in their own right. Though longer eurypterids are known from the eurypterine suborder, notably the largest known arthropod of all time, Jaekelopterus , at 230–260 centimetres (7.5–8.5 ft), hibbertopterids were far bulkier than any of the largest eurypterine eurypterids and as such likely represent the overall heaviest animals in the order.

Description

Restoration of Campylocephalus. CampylocephalusDB117.jpg
Restoration of Campylocephalus .

Hibbertopterids were large mycteropoid eurypterids characterised by their broad prosomas (heads), hastate (e.g. shaped like a gladius , a Roman sword) telsons (which were the posteriormost division of the body) with paired keels and a covering of ornamentation in the shape of scales across their exoskeletons. More shared features are the presence of tongue-shaped scales on the margins of the tergites of the opisthosoma (the abdomen) and that the fourth pair of appendages were covered in spines. The prosoma of the hibbertopterids possessed lobes that were positioned posterolaterally (posteriorly on both sides). [1]

Although Vernonopterus has been estimated to have reached lengths of "just" 50 centimetres (19.7 inches), both Campylocephalus and Hibbertopterus represent true giant eurypterids. The largest specimens of Campylocephalus suggests a size of 140 centimetres (4.6 feet) and at 180–200 centimetres (5.9–6.6 ft), Hibbertopterus scouleri was the largest eurypterid in the entire stylonurine suborder. [2] That hibbertopterids were so broad and compact probably means they were the overall heaviest eurypterids, surpassing longer forms, such as the light-weight pterygotids, in weight. [3]

Compared to other eurypterids, hibbertopterids were highly unusual. Their broad and heavy bodies go against the general eurypterid trend of being streamlined and lightweight and like all other stylonurine eurypterids, they lack swimming paddles. [4] Out of their six pairs of limbs, the second, third and fourth pairs were forward-facing and specialised for gathering food. The leg segments of these limbs were covered with long spines and their ends were covered with sensory organs. The fourth pair of limbs, although also used in feeding, was also used for locomotion, as were pairs five and six. As such, hibbertopterids would have been hexapods (walking on six legs). [5]

Due to the fragmentary nature of both Campylocephalus and Vernonopterus, confidently establishing distinguishing features between the three genera is difficult. Distinguishing Campylocephalus from Hibbertopterus is primarily the fact that the carapace (exoskeleton covering the head) of Campylocephalus is slightly narrower and is broadest at its midsection, rather than at the part which connects to the rest of the body (as it is in Hibbertopterus). [4] The fragmentary Vernonopterus can be distinguished from the others by its distinctive and unique ornamentation. [6]

Preserved cuticle of hibbertopterids is thinner than what is expected of arthropods of their size, though this might be explained by the possibility that only an outer exocuticular region has been preserved in their fossils. [5]

Classification

Restoration of Hibbertopterus. Hibbertopterus scouleri.jpg
Restoration of Hibbertopterus .

The hibbertopterids are classified as part of the superfamily Mycteropoidea within the eurypterid suborder Stylonurina. The family includes the three genera Campylocephalus, Hibbertopterus and Vernonopterus. Up until recently the two genera Cyrtoctenus and Dunsopterus were also recognised as valid and distinct hibbertopterid genera, but the modern consensus is that they represent ontogenetic stages (different developmental stages of the animal throughout its life) of Hibbertopterus. [6] [7] The genus Hastimima , sometimes classified as part of the Hibbertopteridae, [7] has been recovered as part of the Mycteroptidae (the closest relatives of the hibbertopterids) in recent analyses. [4]

The morphology of the hibbertopterids was so unusual that they have been thought to be an order separate to Eurypterida. [8] Recent work however confirms them to be derived members of the suborder Stylonurina, closely related to Drepanopterus and the Mycteroptidae. [1]

The features of genera Campylocephalus and Vernonopterus makes it clear that both genera represent hibbertopterid eurypterids, but the incomplete nature of all fossil specimens referred to them make any further study of the precise phylogenetic relationships within the Hibbertopteridae difficult. Both genera could even represent further synonyms of Hibbertopterus itself, which would make the family monotypic (containing only one immediately subordinate taxon), though the highly incomplete nature of their remains again makes that hypothesis impossible to confirm. [9]

The cladogram below is adapted from Lamsdell (2012), [4] collapsed to only show the superfamily Mycteropoidea.

Mycteropoidea
Drepanopteridae

Drepanopterus pentlandicus

Drepanopterus abonensis

Drepanopterus odontospathus

Mycteroptidae

Woodwardopterus scabrosus

Mycterops mathieui

Hastimima whitei

Megarachne servinei

Hibbertopteridae

Campylocephalus oculatus

Hibbertopterus scouleri

Hibbertopterus wittebergensis

Palaeoecology

The hibbertopterids were derived sweep-feeders, inhabiting freshwater swamps and rivers and feeding by raking through the soft sediment with blades on their anterior appendages to capture small invertebrates. [10]

Sweep-feeding strategies evolved independently in two of the four stylonurine superfamilies, the Stylonuroidea and the Mycteropoidea. In both superfamilies, the adaptations to this lifestyle involves modifications to the spines on their anterior prosomal appendages for raking through the substrate of their habitats. Stylonuroids have fixed spines on appendages II-IV which could have been used as dragnets to rake through the sediments and thus entangling anything in their way. Mycteropoidea show even more extreme adaptations towards a sweep-feeding lifestyle. These adaptations are taken to an even further extreme within the Hibbertopteridae, with appendage IV possessing a blade alongside the appendages II-III (which also have blades in other hibbertopteroids). [1]

The coxae in Hibbertopterus are reduced, leading to part of the food masticatory process being assumed by the laden (plates overlaying the coxae). Some of its species have even further adaptations towards sweep-feeding than other mycteropoids, with its blades modified into comb-like rachis that could entrap smaller prey or other organic food particles. With its coxae being large, it is likely that some species of Hibbertopterus would also feed on relatively large invertebrates when able to. [1]

Related Research Articles

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.

Stylonurina Extinct suborder of arthropods

Stylonurina is one of two suborders of eurypterids, a group of extinct arthropods commonly known as "sea scorpions". Members of the suborder are collectively and informally known as "stylonurine eurypterids" or "stylonurines". They are known from deposits primarily in Europe and North America, but also in Siberia.

<i>Hibbertopterus</i> Extinct genus of arthropods

Hibbertopterus is a genus of eurypterid, a group of extinct aquatic arthropods. Fossils of Hibbertopterus have been discovered in deposits ranging from the Devonian period in Belgium, Scotland and the United States to the Carboniferous period in Scotland, Ireland, the Czech Republic and South Africa. The type species, H. scouleri, was first named as a species of the significantly different Eurypterus by Samuel Hibbert in 1836. The generic name Hibbertopterus, coined more than a century later, combines his name and the Greek word πτερόν (pteron) meaning "wing".

<i>Megarachne</i> Extinct genus of eurypterid

Megarachne is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils of Megarachne have been discovered in deposits of Late Carboniferous age, from the Gzhelian stage, in San Luis, Argentina. The fossils of the single and type species M. servinei have been recovered from deposits that had once been a freshwater environment. The generic name, composed of the Ancient Greek μέγας (megas) meaning "great" and Ancient Greek ἀράχνη (arachne) meaning "spider", translates to "great spider", because the fossil was misidentified as a large prehistoric spider.

<i>Drepanopterus</i> Extinct genus of sea scorpions

Drepanopterus is an extinct genus of eurypterid and the only member of the family Drepanopteridae within the Mycteropoidea superfamily. There are currently three species assigned to the genus. The genus has historically included more species, with nine species associated with the genus Drepanopterus, however five of these have since been proven to be synonyms of pre-existing species, assigned to their own genera, or found to be based on insubstantial fossil data. The holotype of one species proved to be a lithic clast.

<i>Campylocephalus</i> Extinct genus of arthropods

Campylocephalus is a genus of eurypterid, a group of extinct aquatic arthropods. Fossils of Campylocephalus have been discovered in deposits ranging from the Carboniferous period in the Czech Republic to the Permian period of Russia. The generic name is composed of the Greek words καμπύλος (kampýlos), meaning "curved", and κεφαλή (kephalē), meaning "head".

Stylonuridae Extinct family of arthropods

Stylonuridae is a family of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions". The family is one of two families contained in the superfamily Stylonuroidea, which in turn is one of four superfamilies classified as part of the suborder Stylonurina.

Rhenopteridae Extinct family of arthropods

The Rhenopteridae are a family of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions". The family is the only family currently contained in the superfamily Rhenopteroidea, one of four superfamilies classified as part of the suborder Stylonurina.

Mycteroptidae Extinct family of arthropods

Mycteroptidae are a family of eurypterids, a group of extinct chelicerate arthropods commonly known as "sea scorpions". The family is one of three families contained in the superfamily Mycteropoidea, which in turn is one of four superfamilies classified as part of the suborder Stylonurina.

Pterygotioidea Extinct superfamily of eurypterids

Pterygotioidea is a superfamily of eurypterids, an extinct group of aquatic arthropods. Pterygotioids were the most derived members of the infraorder Diploperculata and the sister group of the adelophthalmoid eurypterids. The group includes the basal and small hughmilleriids, the larger and specialized slimonids and the famous pterygotids which were equipped with robust and powerful cheliceral claws.

Stylonuroidea Extinct superfamily of arthropods

Stylonuroidea is an extinct superfamily of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions". It is one of four superfamilies classified as part of the suborder Stylonurina.

Parastylonuridae Extinct family of arthropods

The Parastylonuridae are a family of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions". The family is one of two families contained in the superfamily Stylonuroidea, which in turn is one of four superfamilies classified as part of the suborder Stylonurina.

Kokomopteroidea Extinct superfamily of arthropods

Kokomopteroidea is an extinct superfamily of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions". It is one of four superfamilies classified as part of the suborder Stylonurina. Kokomopteroids have been recovered from deposits of Early Silurian to Late Devonian age in the United States and the United Kingdom.

Kokomopteridae Extinct family of arthropods

The Kokomopteridae are a family of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions". The family is one of two families contained in the superfamily Kokomopteroidea, which in turn is one of four superfamilies classified as part of the suborder Stylonurina.

Hardieopteridae Extinct family of arthropods

The Hardieopteridae are a family of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions". The family is one of two families contained in the superfamily Kokomopteroidea, which in turn is one of four superfamilies classified as part of the suborder Stylonurina. Hardieopterids have been recovered from deposits of Early Silurian to Late Devonian age in the United States and the United Kingdom.

Adelophthalmidae Family of eurypterids, an extinct group of aquatic arthropods

Adelophthalmidae is a family of eurypterids, an extinct group of aquatic arthropods. Adelophthalmidae is the only family classified as part of the superfamily Adelophthalmoidea, which in turn is classified within the infraorder Diploperculata in the suborder Eurypterina.

Eurypterina Suborder of eurypterid

Eurypterina is one of two suborders of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions". Eurypterine eurypterids are sometimes informally known as "swimming eurypterids". They are known from fossil deposits worldwide, though primarily in North America and Europe.

Mycteropoidea Extinct superfamily of arthropods

Mycteropoidea is an extinct superfamily of eurypterids, an extinct group of chelicerate arthropods commonly known as "sea scorpions". It is one of four superfamilies classified as part of the suborder Stylonurina. Mycteropoids have been recovered from Europe, Russia, South America and South Africa. Mycteropoid specimens are often fragmentary, making it difficult to establish relationships between the included taxa. Only two mycteropoid taxa are known from reasonable complete remains, Hibbertopterus scouleri and H. wittebergensis.

<i>Vernonopterus</i> Extinct genus of arthropods

Vernonopterus is a genus of eurypterid, a group of extinct aquatic arthropods. Fossils of Vernonopterus have been discovered in deposits of the Carboniferous period in Scotland. The name of the genus derives from the location where the only known fossil has been discovered, Mount Vernon near Airdrie in Lanarkshire, Scotland. A single species of Vernonopterus is recognized, V. minutisculptus, based on fragmentary fossilized tergites, segments on the upper side of the abdomen. The species name minutisculptus refers to the ornamentation of scales that covers the entirety of the preserved parts of the eurypterid.

Slimonidae Extinct family of eurypterids

Slimonidae is a family of eurypterids, an extinct group of aquatic arthropods. Slimonids were members of the superfamily Pterygotioidea and the family most closely related to the derived pterygotid eurypterids, which are famous for their cheliceral claws and great size. Many characteristics of the Slimonidae, such as their flattened and expanded telsons, support a close relationship between the two groups.

References

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  2. Lamsdell, James C.; Braddy, Simon J. (2009). "Cope's Rule and Romer's theory: patterns of diversity and gigantism in eurypterids and Palaeozoic vertebrates". Biology Letters: rsbl20090700. doi:10.1098/rsbl.2009.0700. ISSN 1744-9561. PMID 19828493. Supplementary information Archived 2018-02-28 at the Wayback Machine
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  4. 1 2 3 4 Lamsdell, James (2012). "Redescription of Drepanopterus pentlandicus Laurie, 1892, the earliest known mycteropoid (Chelicerata: Eurypterida) from the early Silurian (Llandovery) of the Pentland Hills, Scotland". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 103: 77–103. doi:10.1017/S1755691012000072.
  5. 1 2 Jeram, Andrew J.; Selden, Paul A. (1993). "Eurypterids from the Viséan of East Kirkton, West Lothian, Scotland". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 84 (3-4): 301–308. doi:10.1017/S0263593300006118. ISSN   1755-6929.
  6. 1 2 Hughes, Emily Samantha (2019), "Discerning the Diets of Sweep-Feeding Eurypterids Through Analyses of Mesh-Modified Appendage Armature". Graduate Theses, Dissertations, and Problem Reports. 3890.
  7. 1 2 Dunlop, J. A., Penney, D. & Jekel, D. 2018. A summary list of fossil spiders and their relatives. In World Spider Catalog. Natural History Museum Bern
  8. Tollerton, V P (1989). "Morphology, Taxonomy, and Classification of the Order Eurypterida Burmeister, 1843". Journal of Paleontology . 63 (5): 642–657. doi:10.1017/S0022336000041275.
  9. James C. Lamsdell, Simon J. Braddy & O. Erik Tetlie (2010). "The systematics and phylogeny of the Stylonurina (Arthropoda: Chelicerata: Eurypterida)". Journal of Systematic Palaeontology . 8 (1): 49–61. doi:10.1080/14772011003603564.
  10. Selden, P.A., Corronca, J.A. & Hünicken, M.A (2005). "The true identity of the supposed giant fossil spider Megarachne". Biology Letters . 1 (1): 44–48. doi:10.1098/rsbl.2004.0272. PMC   1629066 . PMID   17148124.CS1 maint: multiple names: authors list (link)