Pygidial Spines Research Articles

Funeralaspis n. gen.: a new odontopleurine trilobite from the early Middle Ordovician (Dapingian) of Death Valley, eastern California, USA, and the classification of Ordovician odontopleurines.

Funeralaspis n. gen. (type species F. deathvalleyensis n. sp.) is the oldest known formally named odontopleurine trilobite species, described on the basis of abundant secondarily silicified sclerites of all sclerite types except the rostral plate, from the Dapingian of the Antelope Valley Formation, west flank of the Funeral Range, Death Valley National Park, Inyo County, California. Many Ordovician odontopleurines have been assigned to Diacanthaspis, to the point that it has become something of a taxon of convenience. All of the potential species are listed along with their provenance, preservation, and represented sclerite types. A core group of Diacanthaspis, termed Diacanthaspis (sensu stricto or s.s.) possibly represents a Laurentian clade, consisting of the type species, D. cooperi, and D. lepidus, D. orandensis, D. scitulus, D. secretus, and possibly D. elapsa and D. parvula. This group is characterized by extreme dorsal spinosity, dorsal accessory spines on the base of the genal spine, pygidial spines with lateral fringes of tiny spines, and thoracic and pygidial ring furrows with fringes of tiny spines. Funeralaspis deathvalleyensis has none of the features characteristic of Diacanthaspis and in general scarcely resembles members of this Upper Ordovician group. Remaining species that have been assigned to Diacanthaspis, some of which are very poorly known, should be reclassified as knowledge of Ordovician odontopleurines advances, but for the present are treated as Diacanthaspis (sensu lato or s.l.). A heretofore nearly unknown species described as Ceratocephala maquoketensis from the early Katian of Howard County, Iowa, has not been commented upon in the literature since it was proposed. It is an odontopleurine which in the current state of knowledge should be assigned to the Diacanthaspis (s.l.) group.

The trilobite Pseudoblackwelderia lubrica (Damesellidae) from the Cambrian (Guzhangian) Changhia Formation of Northern Anhui, China, and its potential sexual dimorphism

The North China type Cambrian System is well developed in northern Anhui Province. A large number of specimens of the trilobite Pseudoblackwelderia lubrica were collected from the Guzhangian (later Cambrian) Changhia Formation at Fenghuangshan in northern Anhui, China, prompting a redefinition of the species. Our emended diagnosis includes: a cone-shaped glabella that is broadly to acutely rounded anteriorly; three pairs of glabellar furrows; the preglabellar field being absent or very short; the librigena with a narrow lateral border, a broad general field and long genal spine; the pygidium with four rings and a terminal piece on the axis; six or seven pairs of pygidial spines with the anterior five pairs being short, the posterolateral pair being long and stout, the seventh pair being small or absent. The new specimens suggest that Neimengguasps is a junior synonym of Pseudoblackwelderia. Moreover, the presence or absence of paired spines on the posterior margin of pygidium is attributed to potential sexual dimorphism. Qianping Lei [cicelyapple@126.com], Nature Department, Changzhou Museum, Changzhou, Jiangsu Province, 213022, PR China Qing Liu [qingliu1029@yeah.net], Room 104, Unit A, Building Six, Fenghuanghu No. 1, Xinbei District, Changzhou 213125, PR China Shanchi Peng [scpeng@nigpas.ac.cn], State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing 210008, China; Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, PR China.

Claspers in the mid-CambrianOlenoides serratusindicate horseshoe crab–like mating in trilobites

AbstractSexual reproduction represents a fundamental aspect of animal biology, but the diversity of reproductive strategies among early Paleozoic metazoans remains obscure. Direct evidence of reproductive strategies comes from exceptionally preserved egg masses in Cambrian and Ordovician euarthropods such as waptiids and trilobites, but anatomical or behavioral adaptations for mating in these taxa are all but unknown. We demonstrate the presence of functionally specialized appendages in the mid-Cambrian (Wuliuan) Burgess Shale trilobite Olenoides serratus. New material of O. serratus preserves significantly modified and reduced endopodites underneath the seventh thoracic and first pygidial tergites. Comparison with extant euarthropods indicates that these specialized limbs are functional analogs to claspers, which are used by sexually mature males to grasp females prior to or during mating. The claspers in O. serratus were most likely used by the male to hold onto the posterior pygidial spines of the female during amplex, similar to the strategy observed in the horseshoe crab Limulus polyphemus. The new appendicular data from O. serratus provide the first direct evidence for trilobite sexual dimorphism based on the non-biomineralized ventral anatomy. Our findings illuminate the reproductive biology of trilobites and indicate that complex mating behaviors observed in extant euarthropods originated during the Cambrian explosion.

A revised ontogeny of the early Ordovician trilobite Leptoplastides salteri (Callaway, 1877)

ABSTRACTThe ontogeny of the pelturine olenid trilobite Leptoplastides salteri (Callaway, 1877) from the Shineton Shales, Shropshire, England, was first described in 1925 by Frank Raw. Since that time, scanning electron microscopy and other new technologies have revealed many more details of structure, of early developmental stages in particular, than were available to Raw. Whereas protaspides are not preserved and the state of preservation is less than perfect for the smallest meraspides, we have established that the latter had an array of delicate, long thoracic and pygidial spines, as well as paired procranidial spines, which disappear by meraspid degree 8. Raw's reconstructions of early meraspides, and his measurements of the early stages in development, are here amended in the light of new information. Dorsal spines in the adult are much more highly developed than have been documented in any other olenid. The hypostome is preserved in place in several specimens. Initially conterminant (attached to the doublure), it becomes natant (free) in late meraspid to early holaspid stages of development, with its anterior contour fitting exactly to that of the glabella. The ecology of the widespread Leptoplastides is best known from very extensive sections in South America, which provide a useful basis for comparison. It was well adapted to a range of environments, both oxygenated and dysoxic, and is usually the dominant taxon in the biofacies in which it is found.

The Upper Ordovician trilobiteRaymonditesSinclair, 1944 in North America

AbstractThe Upper Ordovician (Sandbian–Katian) bathyurid trilobiteRaymonditesSinclair is revised using new collections from Missouri and Ontario, and archival material from Illinois, Wisconsin, New York, and Ontario. Phylogenetic analysis supports monophyly ofRaymondites, but recognition of this genus rendersBathyurusBillings paraphyletic. We treatRaymonditesas a subgenus ofBathyurusand label the paraphylum of species traditionally assigned to the latter asBathyurussensu lato.Bathyurus(Raymondites) is composed of five previously named species,B. (R.)spiniger(Hall),B. (R.)longispinus(Walcott),B. (R.)ingalli(Raymond),B. (R.)bandiferSinclair, andB. (R.)trispinosus(Wilson), and two new species,B. (R.)clochensis, andB. (R.)missouriensis; an eighth species is placed in open nomenclature. All species share tuberculate sculpture on the glabella, a relatively short palpebral lobe whose length is less than half of preoccipital glabellar length, and a pygidial outline that is well rounded posteriorly. Aside from the most basal species,B. (R.)longispinus, they also possess occipital spines and, where the pygidium is known, axial pygidial spines.

Late ontogeny of the trilobite Tsinania shanxiensis (Zhang and Wang, 1985) from the Cambrian (Furongian) of Anhui, China and its systematic implications

A new occurrence of Tsinania shanxiensis (Zhang and Wang, 1985) n. comb., is reported herein from northern Anhui, China. This species is here transferred from Guluheia to Tsinania as the differences between the two genera are not enough to support the generic difference. The late ontogeny of T. shanxiensis is documented herein, which provides important evidence for the evolutionary origin of the Family Tsinaniidae. The late ontogeny of the pygidia of T. shanxiensis is divided into four stages: PSS+2 stage, PSS+1 stage, early holaspid stage, and late holaspid stage. The immature morphology of T. shanxiensis retains long pygidial spines, reminicent of Mansuyia. Therefore, the morphology of T. shanxiensis can be regarded as the transitional type between kaolishaniids (e.g., Mansuyia orientalis, M. tani) and tsinaniids (e.g., T. canens, Shergoldia laevigata) during the middle Furongian.

Description and ecology of a new Middle Ordovician (Llanvirn) odontopleurid trilobite from the Builth Inlier of Mid-Wales, with a review of the genusMeadowtownella

AbstractA new odontopleurid trilobite,Meadowtownella serrata, is described from a new Llanvirnian (Darriwilian;Didymograptus murchisoniBiozone) locality in the Builth Inlier, Mid-Wales. This unusually spinose species displays a remarkable array of pygidial spines and extends the stratigraphic range of the genus back into the Abereiddian. Benthic faunas are rare in the predominantly ‘anoxic’D. murchisoniBiozone of the Welsh Basin, and these beds are an unusual habitat for odontopleurids. Environmental analyses (framboidal pyrite and associated total organic carbon) in this study, together with the distribution of benthic faunas (described and figured) in the section, indicate a likely oxic water column and episodically oxic bottom waters. A new diagnosis forMeadowtownellais presented with a review of the genus, as a clarification of previous work. The greater spinosity of earlier members of the lineage may reflect either changing environmental preferences, or the derivation ofMeadowtownellafrom an original more spinose lineage.

Panisaspis, a new genus of pliomerid trilobites from the Lower Ordovician (Ibexian; Tulean and Blackhillsian) of the Great Basin, western USA

Panisaspis n. gen. is a clade of pliomerid trilobites from the Tulean and Blackhillsian stages (Floian) of the Great Basin. It includes Protopliomerops? quattuor Hintze, 1953, and ten new species, six of which are formally named: Panisaspis millardensis (type species), P. sevierensis, P. deltaensis, P. rancherensis, P. topscityensis, and P. loganensis. Four new species are not well enough known for formal naming and are described in open nomenclature. All species are Tulean in age except for P. millardensis, which is earliest Blackhillsian. Synapomorphies of Panisaspis include a short, nearly semicircular anterior border; small L1; large genal spines; a rounded, ovoid hypostomal border; elongated third pygidial spines; and a large, triangular terminal piece with distinct pitted impressions. Phylogenetic analysis indicates that P. millardensis and P. sevierensis are sister taxa, and that P. deltaensis, P. rancherensis, P. topscityensis, P. quattuor, and P. loganensis are successive sister species. The group may be sister to Ibexaspis Přibyl and Vaněk in Přibyl et al., 1985.

Heckethornia, a new genus of dimeropygid trilobites from the Lower Ordovician (Ibexian; Tulean and Blackhillsian) of the Great Basin, western USA

Heckethornia n. gen. is a morphologically striking clade of dimeropygid trilobites known from the Pogonip Group (Ordovician; Ibexian Series, Tulean and Blackhillsian stages) of western Utah and eastern Nevada. It includes seven species based on silicified material. All occur in the Fillmore Formation of western Utah, and four are also known from the Yellow Hill Limestone of eastern Nevada. Diagnostic features of Heckethornia include a highly vaulted exoskeleton with a tall pygidial “wall” made of fused outer pleurae, three pairs of large glabellar spines, two to three pairs of prominent pygidial spines, a single (or paired) large occipital spine(s), and an arc of tubercles on the librigenal field. Cladistic parsimony analysis suggests that the genus comprises two subclades, one including species with a single median occipital spine, and species with paired occipital spines or tubercles in the other. New species are H. smithi , H. hyndeae , H. numani , H. bowiei , H. morrisseyi , and H. ballionae .

Post‐embryonic development of the Furongian (late Cambrian) trilobite Tsinania canens: implications for life mode and phylogeny

The current concept of the order Asaphida was proposed to accommodate some Cambrian and Ordovician trilobite clades that are characterized by the possession of a ventral median suture. The family Tsinaniidae was recently suggested to be a member of the order Asaphida on the basis of its close morphological similarity to Asaphidae. Postembryonic development of the tsinaniid trilobite, Tsinania canens, from the Furongian (late Cambrian) Hwajeol Formation of Korea, reveals that this trilobite had an adult-like protaspis. Notable morphological changes with growth comprise the effacement of dorsal furrows, sudden degeneration of pygidial spines, regression of genal spines, and loss of a triangular rostral plate to form a ventral median suture. Programmed cell death may be responsible for degenerating the pygidial and genal spines during ontogeny. Morphological changes with growth, such as the loss of pygidial spines, modification of pleural tips, and effacement of dorsal furrows, suggest that T. canens changed its life mode during ontogeny from benthic crawling to infaunal. The protaspid morphology and the immature morphology of T. canens retaining genal and pygidial spines suggest that tsinaniids bear a close affinity to leiostegioids of the order Corynexochida. Accordingly, development of a ventral median suture in T. canens demonstrates that the ventral median suture could have evolved polyphyletically, and thus the current concept of the order Asaphida needs to be revised.

Asteropygine trilobites from the Late Devonian of the Kerman region, Iran

The discovery of two new Frasnian species of the subfamily Asteropyginae, Bradocryphaeus kermanensis and Radiopyge hamedii, in the Haruz section north of Kerman, east-central Iran, demonstrates for the first time the presence of North Gondwanan trilobites in this region. The two new species provide a link between faunas from Afghanistan to the east and the Alborz Mountains in north Iran. The recognition of R. hamedii confirms the validity of Radiopyge Farsan, hitherto known only from Afghanistan, as well as the presence in some Frasnian Asteropyginae of six pairs of lateral pygidial spines.

The ontogeny ofTaihungshania miqueli(Bergeron, 1893) from the Arenig of the Montagne Noire, southern France

AbstractThe very large trilobiteTaihungshania miquelioccurs abundantly in siliceous nodules in the Ordovician of the Montagne Noire, southern France, but in these, larval individuals are not preserved. At a new locality at Peret, near Cabrières, protaspid and early meraspid growth stages occur in great abundance, together with a few complete specimens belonging to meraspid degrees 1, 2, 3, 5 and 7, and a single young holaspis. Although this material is largely flattened, and the originally globular protaspides crushed, it is still possible to establish the development of several axial rings before the transitory pygidium becomes liberated. In the degree 0 meraspis, the cephalon and transitory pygidium are about the same size, and the presence of a partially enrolled degree 3 meraspis indicates that protective coiling was possible by this early stage. Palpebral lobes are incipient in degree 2 meraspides, and fully formed by degree 7, otherwise the morphology of the cephalon becomes quite adult-like by degree 4. The pygidial spines begin to appear in degree 5, at which stage there are about 10 axial rings; there are about 13 in a degree 7 meraspis, and in the adult, where the posterior contour is semi-elliptical, there are some 22. The smallest hypostomes present belong to a degree 6 meraspis, and these are morphologically similar to those of adults. Observations on the later growth stages were made on material from another locality (Caramaou), as well as that from Peret. In the eye the upper lenses are large and slightly separated, but those towards the base are much diminished in size; this is a function of an unusual growth geometry.At Peret, all stages of development, at least as far as young adults, are represented, albeit often in a fragmentary state. As with other asaphines, the protaspides are assumed to be planktic, andT. miquelipresumably left the plankton and became benthic between the protaspid and meraspid periods.

New Early Devonian styginid trilobites from Victoria, Australia, with revision of some spinose styginids

A distinctive new genus (Xyoeax) and two new species (Xyoeax eponcus, Scutellum sudorum) of styginid trilobites are described from the late Early Devonian (Emsian) Murrindal Limestone of eastern Victoria, Australia. Limited material of a third styginid species not belonging to any named genus is also described from the same locality. Xyoeax is considered to be closely related to Ancyropyge Clarke, 1891, and Weberopeltis Maksimova in Moore, 1959, and less closely related to Kolihapeltis Prantl and Přibyl, 1947. Weberopeltis is revised and newly diagnosed. Several species previously assigned to Weberopeltis are transferred to the new genus Exastipyx, which is considered to be related to Cornuscutellum Snajdr, 1960. Ancyropyge, Exastipyx, Weberopeltis, and Xyoeax all have pygidia with long pleural spines derived by reduction of the pleural field between the pleural ribs, whereas Kolihapeltis and Cornuscutellum lack pygidial spines. Pygidial morphology of Xyoeax type has thus arisen independently in separate styginid lineages. The type species of Altaepeltis Maksimova, 1968 is established as Weberopeltis bublitchenkoi Maksimova, 1960, by monotypy.

Spine function in the odontopleurid trilobites Leonaspis and Dicranurus from the Devonian of Oklahoma

The Lower Devonian Haragan Formation of Oklahoma has well preserved odontopleurid trilobites found in repeated calcarous mudstones. These layers (5–9cm. thick) represent distal tempestites that rapidly buried and preserved completely articulated odontopleurids which provide important paleobiological information. The genal and pleural spines of Leonaspis williamsi and Dicranurus hamatus elegantus of the Haragan Formation are examined and compared.In Leonaspis the ventrally directed genal and pleural spines are straight and lay flat. The first pleural spine is the longest followed by progressively shorter pleural spines. Several collected and prepared enrolled specimens have the spines radiating outward. In an enrolled position the spines would protect Leonaspis from a large predator.The pleural spines of Dicranurus have a very different pattern from Leonaspis. The first three spines are short and laterally directed, the fourth, sixth and seventh are very long, curved and directed ventrally, the eighth, ninth and pygidial spines are short and decrease in length toward the pygidium. There is no pleural spine on the fifth thoracic segment and the genal spine is very long and curved. On these spines at many growth stages there is direct and indirect evidence of being encrusted based on the following: (1) Epibionts such as bryozoans, brachiopods and a crinoid; 2) Endoliths (25–150 microns in diameter) of unknown origin are found all over Dicranurus but are predominantly on the spines; 3) Pyrite (now limonite) coating the spines apparently formed from the decay of organic matter. If the source of the organic matter came from the trilobite, then pyrite would be found all over Dicranurus. However, it is only found randomly on the spines. This pattern is like some of the encrusting bryozoans suggesting that the source of the organic matter may be from an encrusting organism.It is doubtful that the pleural spines of Dicranurus would be adapted for resting on the bottom when bryozoans are found encrusting the entire surface of the spine. However, the very long pleural and genal spines with attched organisms would break up the outline of Dicraurus making it less conspicuous to a predator.

A FUNCTIONAL STUDY OF THE SILURIAN ODONTOPLEURID TRILOBITE LEONASPIS DEFLEXA (LAKE)

The odontopleurid trilobite Leonaspis deflexa (Lake) was functionally capable of life in two alternative attitudes. In the ‘resting’ attitude the whole body could lie outstretched on the sea floor, supported on the hindmost anterior denticles of the cephalon and on the genal, thoracic, and pygidial spines. In this attitude the body declined posteriorly. In the ‘active’ attitude, however, the cephalon was tilted forwards through 25°, bringing the body up horizontally. The hypostome and presumably the mouth also could then have been brought into close proximity with the sea floor. Leonaspis, Primaspis, Diacanthaspis, and Anacaenaspis are all functionally comparable. Dudleyaspis, Acidaspis, and other genera were specialised permanently for life in an attitude analogous to the ‘active’ posture of Leonaspis. The life attitudes postulated for odontopleurids are compared with those of other trilobites and some comments are given on the evolution of the different patterns.