THE SPECIES GROUPS OF GONATOCERUS NEES IN NORTH AMERICA WITH A REVISION OF THE SULPHURIPES AND ATER GROUPS (HYMENOPTERA: MYMARIDAE)

1988 ◽  
Vol 120 (S141) ◽  
pp. 3-110 ◽  
Author(s):  
John T. Huber

AbstractA provisional cladistic analysis of the species groups of Gonatocerus Nees is presented, based on 24 characters. Six species groups are defined, but only the four Nearctic ones are described. The species groups proposed are the ater, deficiens, litoralis, membraciphagus, straeleni, and sulphuripes groups. The ater group is divided into the morrilli, ater (s.s.), and bucculentus subgroups. Keys are provided to distinguish the species groups, and females of the sulphuripes- and ater-group species. Twenty-four species are recognized in the sulphuripes and ater groups for America north of Mexico. Eight species are recognized in the sulphuripes group, of which californicus, koebelei, maga, mexicanus, rivalis, and utahensis are redescribed. Two species, floridensis and lissonotus, are described as new. One new synonymy is proposed: titillatus is considered a junior synonym of utahensis. Sixteen species are recognized in the ater group, of which ashmeadi, capitatus, dolichocerus, fasciatus, latipennis, morrilli, novifasciatus, ovicenatus, and triguttatus are redescribed. Seven species, bucculentus, enicmophilus, flagellatus, impar, incomptus, inexpectatus, and inflatiscapus, are described as new. Two new synonymies are proposed: maximus is considered a junior synonym of latipennis, and marilandicus a junior synonym of dolichocerus. Lectotypes are designated for ashmeadi, capitatus, koebelei, mexicanus, morrilli, ovicenatus, triguttatus, and utahensis.

1993 ◽  
Vol 125 (S168) ◽  
pp. 1-193 ◽  
Author(s):  
Valerie M. Behan-Pelletier

AbstractThe oribatid family Eremaeidae is represented in North America by two genera, Eremaeus and Eueremaeus, both widely distributed throughout the Palaearctic and Nearctic regions. In North America species in both genera are found in moist to arid habitats from New Mexico to the High Arctic. Reproduction is sexual, and both immatures and adults feed mainly on fungi.Revised diagnoses are presented for the Eremaeidae and genera Eremaeus and Eueremaeus. Eighteen species of Eremaeus, of which 14 are newly proposed, and 24 species of Eueremaeus, of which 15 are newly proposed, are recognized. Identification keys are provided for the world genera of Eremaeidae, and for adults of Eremaeus and Eueremaeus of North America. All but one North American species of these genera are described, and their geographical distributions mapped.North American Eremaeus species include E. appalachicus sp. no v., E. boreomontanus sp. nov., E. brevitarsus (Ewing), E. californiensis sp. nov., E. gracilis sp. nov., E. grandis Hammer, E. kananaskis sp. nov., E. kevani sp. nov., E. megistos sp. nov., E. monticolus sp. nov., E. nortoni sp. nov., E. occidentalis sp. nov., E. oresbios sp. nov., E. plumosus Woolley, E. porosus sp. nov., E. salish sp. nov., E. translamellatus Hammer, and E. walteri sp. nov. The immatures of four of these, E. kananaskis, E. occidentalis, E. oresbios, and E. translamellatus, are described.North American Eueremaeus include Eu. acostulatus sp. nov., Eu. aridulus sp. nov., Eu. columbianus (Berlese), Eu. foveolatus (Hammer), Eu. marshalli sp. nov., Eu. masinasin sp. nov., Eu. michaeli sp. nov., Eu. nahani sp. nov., Eu. nemoralis sp. nov., Eu. proximus (Berlese) comb, nov., Eu. woolleyi (Higgins) comb, nov., Eu. yukonensis sp. nov., and three informal species groups with the following included species in North America: (1) Eu. trionus group—Eu. trionus (Higgins) comb, nov., (2) Eu. stiktos group—Eu. carinatus sp. nov., Eu. higginsi sp. nov., Eu. stiktos (Higgins) comb, nov., Eu. tetrosus (Higgins) comb, nov., (3) Eu. chiatous group—Eu. alvordensis sp. nov., Eu. aysineep sp. nov., Eu. chiatous (Higgins) comb, nov., Eu. danos sp. nov., Eu. lindquisti sp. nov., Eu. magniporosus (Wallwork) comb, nov., and Eu. osoyoosensis sp. nov. The immatures of nine of these, Eu. masinasin, Eu. nahani, Eu. carinatus, Eu. higginsi, Eu. columbianus, Eu. proximus, Eu. woolleyi, Eu. stiktos, and Eu. tetrosus, are described. Kartoeremaeus reevesi Higgins and Eremaeus politus Banks are considered junior subjective synonyms of Eueremaeus columbianus (Berlese).A cladistic analysis of the genera comprising Eremaeidae: Eremaeus, Tricheremaeus, Eueremaeus (and included species groups), Proteremaeus, Carinabella, and Asperemaeus, indicates that Eremaeus is the sister taxon of Carinabella, and that Eueremaeus is the sister taxon of Proteremaeus. Tricheremaeus is the sister taxon of Eremaeus + Carinabella, and Asperemaeus is the sister taxon of Eueremaeus + Proteremaeus. The relationship of the Eremaeidae to the Megeremaeidae and Zetorchestidae is presented. Finally, I discuss the ecology and distribution of North American species of Eremaeidae.


1980 ◽  
Vol 112 (S114) ◽  
pp. 5-167 ◽  
Author(s):  
Gary A. P. Gibson

AbstractThe species of the genus Macrophya Dahlbom (Tenthredininae) of North America are revised. Forty-six species and two subgenera are recognized, Macrophya s. str. with 45 species, and Pseudomacrophya Enslin with a single species. The subgenus Paramacrophya Forsius is newly synonymized with Macrophya s. str. Eleven species are described as new: amediata, epinolineata, macgillivrayi, masneri, masoni, mensa, nirvana, phylacida, senacca, serratalineata, and smithi; and the following new synonymy proposed: albomaculata (Norton) = contaminata Provancher, errans Rohwer, and fuscoterminata Rohwer; bifasciata (Say) = pulcherrima Kirby; cassandra Kirby = albilabris Harrington, bellula MacGillivray, and externiformis Rohwer; flavolineata (Norton) = confusa MacGillivray, crassicornis Provancher, (?) lineata Norton, and proximata Norton; flicta MacGillivray = fistula MacGillivray; fuliginea Norton = castanea Rohwer; fumator Norton = pumila Norton, and subviolacea Cresson; mixta MacGillivray = bilineata Dyar; nigra (Norton) = minuta MacGillivray; oregona Cresson = obaerata MacGillivray, and oregona var. dukiae Ross; pannosa (Say) = flaccida MacGillivray, and raui Rohwer; pulchella (Klug) = albifacies Kirby, dyari Rohwer, ornata MacGillivray, and trosula (Norton); pulchelliformis Rohwer = sambuci Rohwer; trisyllaba (Norton) = zonalis Norton; and varia (Norton) = fascialis Norton, and fascialis var. puella Ross. Macrophya varius var. festana Ross is removed from synonymy with M. varia (Norton) and accorded specific status, and the following species removed from previous synonymy and recognized as valid species: simillima Rohwer from intermedia (Norton); flavolineata (Norton), lineatana Rohwer, and zoe Kirby from pulchella (Klug); and oregona Cresson from pumila Norton. Macrophya dejecta (Norton) is removed from the list of Nearctic species, and is removed from synonymy with M. cinctula (Norton) but synonymized with the Palearctic species M. ligata (Müller). Macrophya annulipes Cresson and M. coquilletti (Rohwer) are considered to belong to an undescribed genus and are excluded from Macrophya; and Allantus cestus Say, and the fossiliferous species Macrophya pervetusta Brues, are considered as unrecognized taxa of Tenthredinidae. Neotypes are designated for Allantus intermedius Norton and A. epinotus Say, and lectotypes designated where necessary. The limits of the genus and subgenera are discussed, and the Palearctic and Nearctic faunas compared through species-groups. Keys to the males and females are given, all species described, and the taxonomic characters used to separate the Nearctic species illustrated in figures taken by electron photography and microphotography.


1989 ◽  
Vol 121 (S148) ◽  
pp. 5-168 ◽  
Author(s):  
Jeffrey M. Cumming

AbstractThe genus Symmorphus Wesmael is distributed throughout the Oriental, Palearctic, and Nearctic regions, as well as the northernmost Neotropical region, and appears to occupy a relatively isolated phylogenetic position within the Eumeninae. Females of Symmorphus, for those species in which nesting behavior is known, mass-provision cells in pre-existing cavities with chrysomelid, curculionid, or microlepidopterous larvae.Thirty-six species are recognized, including three new species and two new subspecies. Keys are provided for the identification of adult specimens, all species are described, and their geographical distributions mapped.Two subgenera, S. (Parasymmorphus) Cumming and Vecht and S. (Symmorphus) Wesmael, are recognized. Symmorphus (Parasymmorphus) includes S. parvilineatus (Cameron), S. momunganensis (Schulthess), and S. negrosensis Cumming and Vecht. Within S. (Symmorphus) three informal species groups are recognized, with the following included species and subspecies: (1) S. hoozanensis group — S. hoozanensis (Schulthess), S. alkimus alkimus Cumming and Vecht, S. alkimus dialeukus ssp.nov. (type-locality Sibolangit, N. Sumatra), S. ambotretus sp.nov. (type-locality Mt. Omei, Sichuan Province, China), and S. canlaonicus sp.nov. (type-locality Mt. Canlaon, Negros Oriental, Philippines); (2) S. declivis group — S. declivis Harttig; (3) S. murarius group — S. murarius (Linnaeus), S. foveolatus Gussakovskij, S. glasunowi Morawitz, S. sichuanensis Lee, S. crassicornis (Panzer), S. captivus (Smith), S. angustatus (Zetterstedt), S. albomarginatus albomarginatus (Saussure), S. albomarginatus midas ssp.nov. (type-locality Bentsen Rio Grande Valley State Park, Texas, USA), S. decens (Kostylev), S. violaceipennis Soika, S. mizuhonis Tsuneki, S. bifasciatus (Linnaeus), S. debilitatus (Saussure), S. cliens Soika, S. connexus (Curtis), S. lucens (Kostylev), S. sublaevis (Kostylev), S. canadensis (Saussure), S. projectus Bohart, S. tukvarensis (Meade-Waldo), S. nipteroides sp.nov. (type-locality N. Yourdou, India), S. fuscipes (Herrich-Schaeffer), S. apiciornatus (Cameron), S. aurantiopictus Soika, S. allobrogus (Saussure), S. cristatus (Saussure), S. gracilis (Brullé), and S. paralleliventris Soika.The following new synonymy is proposed: S. calciatii (Zavattari) = S. glasunowi Morawitz; S. nursei (Bingham), S. hospes (Nurse), and S. hospitans (Dalla Torre) = S. crassicornis (Panzer); S. hakutozanus Tsuneki and S. nansetsurei Tsuneki = S. angustatus (Zetterstedt); S. walshianus (Saussure) and S. meridionalis Viereck = S. albomarginatus albomarginatus (Saussure); S. kurentzovi Kurzenko, S. iiyamai Tsuneki, S. shiroyamai Tsuneki, S. piceanus Tsuneki, and S. sassai Tsuneki = S. mizuhonis Tsuneki; S. sinuatus (Fabricius), S. sinuatus mutinensis (Baldini), S. sparsus Morawitz, S. sinuatissimus Richards, S. mutinensis auster Soika, and S. mutinensis yezoanus Tsuneki = S. bifasciatus (Linnaeus); S. ishikawai Soika = S. lucens (Kostylev); S. seoulensis Tsuneki = S. apiciornatus (Cameron); S. pumilus (Saussure) and S. trisulcatus Cameron = S. cristatus (Saussure); S. elegans libanicus Soika = S. gracilis (Brullé); and S. allobrogus (Saussure) is removed from synonymy with S. bifasciatus (Linnaeus). Odynerus latipennis Smith and Odynerus cilicius Cameron, previously considered to belong within Symmorphus, are excluded from the genus, and S. sounkionis Tsuneki is transferred as Ancistrocerus sounkionis (Tsuneki) comb.nov. Neotypes are designated for S. crassicornis (Panzer) and S. fuscipes (Herrich-Schaeffer), and lectotypes are designated for S. foveolatus Gussakovskij, S. glasunowi Morawitz, S. nursei (Bingham), S. laeviventris (Thomson), S. albomarginatus (Saussure), S. decens (Kostylev), S. sinuatus (Fabricius), S. sinuatus mutinensis (Baldini), S. debilitatus (Saussure), S. lucens (Kostylev), S. sublaevis (Kostylev), S. debilis (Saussure), S. apiciornatus (Cameron), S. philadelphiae (Saussure), and S. elegans (Wesmael).A cladistic analysis of Symmorphus, based on 26 characters, is presented, including a discussion of the relationships of the genus to the remainder of the Eumeninae. Hypotheses outlining the historical zoogeography of Symmorphus are presented in light of current paleogeographic and paleoclimatic reconstructions, by comparing these cladistic relationships with the chorological patterns observed within the genus and other Vespidae.


Zootaxa ◽  
2011 ◽  
Vol 2829 (1) ◽  
pp. 1 ◽  
Author(s):  
DAVID S. SELDON ◽  
RICHARD A. B. LESCHEN

The flightless carabid genus Mecodema Blanchard 1843 is restricted to New Zealand and presently contains 64 species. In this study, we examine species in the newly constituted curvidens group (sulcatum and curvidens are synonymised), reducing the Mecodema species groups to seven. They share two synapomorphies (rounded apical lobe of the aedeagus and lack of microsculpture on the vertex of the head) and are distributed in Northland, south along the east coast of the North Island and the northeast portion of the South Island. Adult specimens of the curvidens and sulcatum groups, along with exemplars of the other Mecodema groups (alternans, costellum, ducale, infimate, laterale and spiniferum), and an outgroup Oregus Putzeys 1868 were examined for cladistic analysis using a data matrix composed of a 63 characters and 21 terminal taxa. The analysis resulted in 18 most parsimonious trees. The following new species in the curvidens group are described: M. aoteanoho sp. n., M. haunoho sp. n., M. manaia sp. n., M. parataiko sp. n., M. ponaiti sp. n., and M. tenaki sp. n. Mecodema exitiosus was wrongly sysnonymised under M. occiputale and we propose a new synonymy M. curvidens (=M. exitiosus). We consider the parts of the male genitalia in detail with special attention to the structures of apical plate once everted from the endophallus.


1991 ◽  
Vol 123 (S154) ◽  
pp. 3-104 ◽  
Author(s):  
Darren A. Pollock

AbstractThe classification of the nine world species of Pytho Latreille is reviewed by study of adult, larval, and pupal stages. Keys are provided for separation of species in these three life stages. Taxonomic changes (senior synonym in brackets) include synonymy of P. fallax Seidlitz 1916 [= P. niger Kirby 1837]; P. americanus Kirby 1837 [= P. planus (Olivier 1795)]; P. deplanatus Mannerheim 1843 is transferred from a junior subjective synonym of P. depressus (Linnaeus 1767) to a junior subjective synonym of P. planus (Olivier 1795). Lectotype designations are provided for the following: P. seidlitzi Blair 1925; P. nivalis Lewis 1888; P. niger Kirby 1837; P. fallax Seidlitz 1916; P. abieticola J. Sahlberg 1875; and P. americanus Kirby 1837. Eight larval stage, and 12 adult stage characters were selected for cladistic analysis. Lacking out-group material, pupal characters were not analysed. Character states were polarized using a generalized out-group composed of the three other genera of Pythinae (all monobasic). Phylogenetic analysis based on these 18 characters suggests four monophyletic species-groups: P. seidlitzi group (P. seidlitzi Blair — North America); P. kolwensis group (P. strictus LeConte – North America, P. kolwensis C. Sahlberg —Fennoscandia and the U.S.S.R., P. nivalis Lewis — Japan); P. niger group (P. niger Kirby — North America, P. abieticola J. Sahlberg — Europe, P. jezoensis Kôno — Japan); P. depressus group [P. planus (Olivier, 1795) — North America, P. depressus (Linnaeus, 1767) — Europe and the U.S.S.R.]. Larval stage synapomorphies are relatively more important in defining the species-groups than are those of the adult stage. The ancestor of Pythidae may have been associated with Coniferae as early as the Jurassic. The common ancestor of Northern Hemisphere Pythinae became isolated upon Laurasia once separation from Gondwanaland occurred near the end of the Jurassic. Two of the species-groups have similar disjunctions in North America, Europe, and Japan. The relatively eastern distributions of the North American member of each suggests that the ancestor of each species-group was Euramerican, and underwent vicariance with the opening of the North Atlantic in the Middle Cretaceous. The present distribution of both species-groups is thought to have been caused by the same vicariant event. The ancestor of the P. depressus group, which is presently circumboreal, was probably widespread and could have been Asiamerican in distribution. In the middle to late Tertiary, evidence suggests that Beringia was covered with coniferous forest, and the ancestor of the P. depressus group probably extended across this land bridge. Final separation between any North American and European/Asian species occurred in the Late Miocene or Pliocene, when a cooling climate made possible the evolution of treeless tundra in the north.


Zootaxa ◽  
2003 ◽  
Vol 276 (1) ◽  
pp. 1 ◽  
Author(s):  
XUEKUI SUN ◽  
STEPHEN A. MARSHALL

The genus Phasia (Diptera, Tachinidae) is redefined and the world species (excluding Neotropical species) are revised. Six species-groups are defined, and seventy-five species are described or redescribed, including 31 new species. Revised host lists and distribution maps are given. Fifteen generic names are treated as new synonyms of Phasia: Akosempomyia Villeneuve, Alophorophasia Townsend, Androeuryops Beneway, Besserioides Curran, Bogosiella Villeneuve, Campbellia Miller, Efftayloria Malloch, Epaulophasia Townsend, Heyneophasia Townsend, Kosempomyia Villeneuve, Paraphasiana Townsend, Tayloria Malloch, Trichophasia Townsend, Xanthotrichius Townsend and Xiphophasia Townsend. The following new species are described: Phasia africana, P. australiensis, P. brachyptera, P. cana, P. clavigralla, P. bifurca, P. cylindrata, P. distincta, P. faceta, P. frontata, P. furcata, P. japanensis, P. kuodi, P. lauta, P. malaisei, P. malayana, P. mathisi, P. minima, P. nirgromaculata, P. rotundata, P. serrata, P. siberica, P. sichuanensis, P. singuliseta, P. subnitida, P. sumatrana, P. transvaalensis, P. triangulata, P. wangi, P. woodi and P. yunnanica. The following new synonyms are proposed: P. umbrifera (Wulp), P. occidentalis (Brooks), P. opaca (Coquillett), P. polita (Brooks), P. alaskensis (Brooks) and P. umbrosa (Wulp) are junior synonyms of P. aeneoventris (Williston); P. nigra (Brooks) is a junior synonym of P. albipennis (Brooks); P. karczewskii (Draber-Mo½ko) is a junior synonym of P. aldrichii (Townsend); P. splendida (Coquillett) and P. meliceris (Reinhard) are junior synonyms of P. aurulans Meigen; P. cockaynei (Miller) and P. lancifer (Malloch) are junior synonyms of P. campbelli (Miller); P. peruviana (Townsend), P. pollinosa (Brooks), P. auricaudata (Brooks) and P. dimidiata (Townsend) are junior synonyms of P. chilensis (Macquart); P. basalis (Malloch), P. hyalis (Malloch) and P. nigrihirta (Malloch) are junior synonyms of P. lepidofera (Malloch); P. bancrofti (Paramonov) is a junior synonym of P. normalis (Curran); P. celer (Townsend) and P. argentifrons (Brooks) are junior synonyms of P. purpurascens (Townsend); P. fumosa (Coquillett), P. nitida (Coquillett), P. phasioides (Coquillett), P. pulverea (Coquillett), P. furva West and P. brevineura West are junior synonyms of P. robertsonii (Townsend); P. chrysis (Malloch), P. aureiventris (Curran), P. costalis (Malloch), P. nigrisquama (Malloch) and P. discalis (Malloch) are junior synonyms of P. rufiventris (Macquart); P. lativentris (Malloch) is a junior synonym of P. sensua (Curran). A cladistic analysis of Phasia, based on fifty-one characters, is presented and the zoogeography of the genus is discussed.


1996 ◽  
Vol 128 (1) ◽  
pp. 85-114 ◽  
Author(s):  
L. Masner ◽  
J. Denis

AbstractThe state of taxonomy of Idris, egg parasitoids of spiders, in Nearctic North America is reviewed. The concept of Idris is discussed and Ceralobaeus Ashmead is considered a junior synonym (new synonymy). The melleus-group of Idris is defined and 11 Nearctic species are keyed and described: I. castaneus sp.nov. (south and southeastern USA), I. chrysion sp.nov. (southeastern USA), I. costatus sp.nov. (eastern USA), I. lacunatus (Baja California, Mexico). I. leedsi sp.nov. (midsouthem USA), I. melleus (Ashrnead) (south and southeastern USA and southern Canada), I. onychion sp.nov. (southeastern USA), I. ornatus sp.nov. (eastern USA), I. pulvinus sp.nov. (south and southeastern USA), I. spartinae sp.nov. (southeastern USA), and I. triticola sp.nov. (midwestern USA). Extra-limital Nearctic members of the melleus-group are briefly discussed.


2017 ◽  
Vol 31 (4) ◽  
pp. 442 ◽  
Author(s):  
Miguel Machado ◽  
Renato A. Teixeira ◽  
Arno A. Lise

The present paper presents a cladistic analysis of the spider genus Tobias Simon, 1895. The analyses were based on a matrix with 29 terminal taxa scored for 86 morphological characters, with a dataset tested under two methodologies for character weighting (i.e. equal and implied weighting). Both analyses supported the paraphyletic relationship of Tobias with Epicadus Simon, 1895, with the former considered a junior synonym of the latter (new synonymy). Onocolus, Epicadinus and Epicadus form a clade supported by two synapomorphies, corroborating previous taxonomic assumptions regarding the proximity of these genera. Epicadus has two lineages: the ‘pustulosus clade’, comprising bark-like individuals that camouflage themselves on tree trunks and branches, and the ‘heterogaster clade’, which consists of a group of spiders that present a wide range of polychromatism and use flowers as hunting sites. Synonymies are presented and a new combination for all species previously assigned as part of Tobias is proposed. Epicadus now comprises nine species.


2002 ◽  
Vol 33 (1) ◽  
pp. 91-102 ◽  
Author(s):  
Masanori J. Toda ◽  
Yao-Guang Hu

AbstractPhylogenetic relationships among the genus Dichaetophora, the genus Nesiodrosophila and the Lordiphosa tenuicauda species-group and some possibly related genera of Drosophilinae were analyzed in this study using 30 morphological characters derived from 34 species. It is concluded that the three taxa constitute a monophyletic group, within which three monophyletic groups are recognized: Dichaetophora + Nesiodrosophila comprise a monophyletic group, while the L. tenuicauda group is divided into two monophyletic groups. This clade is revised as the genus Dichaetophora, with its three constituent monophyletic groups treated as new specis-groups: the agbo, the tenuicauda and the acutissima groups. Nesiodrosophila becomes a junior synonym. A key to the species-groups is provided.


1990 ◽  
Vol 64 (4) ◽  
pp. 600-614 ◽  
Author(s):  
Jonathan M. Adrain ◽  
Brian D. E. Chatterton

Odontopleura (Odontopleura) arctica, a new species of odontopleurine trilobite, is described from the Canadian Arctic. A method of cladistic analysis is detailed. Parsimony analysis should be performed treating all characters as unordered. The universe of directed trees implied by the resulting rootless network(s) can then be examined and a preferred tree selected by a criterion of congruency. Namely, the most parsimonious directed tree that accommodates the most congruent arrangement of character-states should be taken as the preferred cladogram. Since this is essentially a general congruency method operating within the constraints of parsimony, it is termed “constrained congruency.” The method is applied to the genus Odontopleura, resulting in the recognition of two major species groups, the nominate subgenus and Sinespinaspis n. subgen. Odontopleura (Ivanopleura) dufrenoyi Barrande is tentatively included in the genus, but considered too poorly known for cladistic analysis. Species assigned to Odontopleura (Odontopleura) include Odontopleura ovata Emmrich, Odontopleura brevigena Chatterton and Perry, Odontopleura (Odontopleura) arctica n. sp., and Diacanthaspis serotina Apollonov. Species assigned to Sinespinaspis n. subgen. include Taemasaspis llandoveryana Šnajdr, Odontopleura greenwoodi Chatterton and Perry, Odontopleura maccallai Chatterton and Perry, and Odontopleura nehedensis Chatterton and Perry. Odontopleura bombini Chatterton and Perry is tentatively placed in synonymy with Odontopleura nehedensis. The genus had a wide distribution throughout the Early and Middle Silurian, due to preferences for deep-water, distal shelf or shelf-slope transition zone habitats.


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