Topotypic Material Research Articles

Next-Generation Sequencing of an 88-Year-Old Specimen of the Poorly Known Species Liagora japonica (Nemaliales, Rhodophyta) Supports the Recognition of Otohimella gen. nov.

Liagora japonica is a red algal species distributed in temperate regions of Japan. This species has not been collected from its type locality on the Pacific coast of Japan since 1927 and seems to have become extinct in this area. For molecular characterization of L. japonica, we extracted DNA from the topotype material of L. japonica collected in 1927, analyzed seven genes using Illumina next-generation sequencing, and compared these data with sequences from modern samples of similar red algae collected from the Japan Sea coast of Japan. Both morphological and molecular data from modern samples and historical specimens (including the lectotype and topotype) suggest that the specimens from the Pacific and Japan Sea coasts of Japan should be treated as a single species, and that L. japonica is phylogenetically separated from the genus Liagora. Based on the phylogenetic results and examination of reproductive structures, we propose Otohimella japonica gen. et comb. nov., characterized morphologically by diffuse carposporophytes, undivided carposporangia, and involucral filaments initiated only from the cortical cell on the supporting cell.

Morphometric Data on Topotype Assemblage of Miogypsina (Lepidosemicyclina) droogeri, Foraminifera from Kachchh, Gujarat

Abstract: Miogypsina (Lepidosemicyclina) droogeri Zone occupies an important position in the Indian biochronostratigraphy since the reservoirs of the giant oil field of Bombay High pertain to this zone. A rich assemblage of the topotype material from Kachchh is morphometically analysed for the better understanding of the variation in the population and its relation with the closely related species. The mean value of V is recorded as 77.5 for 24 specimens of Miogypsina (Lepidosemicyclina) droogeri Mohan and Tewari. The detailed study of Miogypsinidae from the exposures in Kachchh and a continuously cored offshore well ED-D from Bombay Offshore has helped in the better understanding of the evolution of Lepidosemicyclina lineage from Miogypsina (Lepidosemicyclina) thecideaeformis Rutten to Miogypsina (Lepidosemicyclina) droogeri Mohan and Tewari through Miogypsina (Lepidosemicyclina) talukdari Raju and Singh.

Morphological distinction between estuarine polychaetes: Laeonereis culveri and L. nota (Phyllodocida: Nereididae).

The family Nereididae includes more than 500 polychaete species described worldwide, and includes species common in many benthic environments, but some other species may tolerate freshwater or can even thrive in humid substrates in tropical forests. In estuarine environments, nereidid polychaetes can be abundant and relevant as a food source for resident or migratory birds. Laeonereis culveri (Webster, 1879) is a common estuarine species found in tropical and subtropical Atlantic American shores and was described from New Jersey; its median and posterior parapodia have upper notopodial ligules usually longer than the lower ones, and the latter are parallel to the notaciculae throughout the body. L. culveri distribution is from Connecticut to central Argentina; however, this wide distribution might be due to the inclusion of several other species as junior synonyms, despite that some morphological differences were found between them. One of such species is L. nota (Treadwell, 1941), that was described from Texas; its parapodia have notopodial ligules of about the same size, and the lower ones are oblique to the notaciculae. In order to clarify the differences between these two species, and to define which inhabits the Northwestern Caribbean region, topotype materials from these two species and specimens from Chetumal Bay were collected, and their morphological features were compared. Our results indicated that L. culveri and L. nota are different species and that the latter is found in Chetumal Bay. On the basis of mature specimens, L. culveri is hereby restricted to the Northern Gulf of Mexico and Northwestern Atlantic Ocean, and L. nota are reinstated and its distribution extends from Texas, in the Gulf of Mexico to Chetumal Bay, in the Northwestern Caribbean Sea. A key to identify all species in Laeonereis Hartman (1945) is also included.

A new reconstruction of continentalTreptichnusbased on exceptionally preserved material from the Jurassic of Massachusetts

AbstractContinental deposits of the Early Jurassic East Berlin Formation in Holyoke, Massachusetts, have yielded an exceptional occurrence of the ichnogenusTreptichnus. Here, burrows are preserved in full relief within thin mud laminae between layers of fine-grained, cross-bedded sandstone. We studied these burrows to evaluate whether earlier explanations of burrow morphology are applicable to allTreptichnus. Our research focused on three questions. (1) Do the HolyokeTreptichnushave significant vertical relief? (2) Does the lack of projections in some of the HolyokeTreptichnusresult from stratinomic sectioning through the bottom of the burrow? (3) Do expanded, bulbous ends of burrow segments result from sediment compaction? While addressing these questions, the Holyoke fossils were compared to syntype and topotype material ofTreptichnusfrom the Carboniferous of Indiana. The HolyokeTreptichnusdid not exhibit significant vertical relief, and the presence and absence of projections is explained by the positioning of new segments at different points along older ones. The bulbous ends of burrow segments resulted from trace-maker behavior, not sediment compaction. Drawing on the analysis of the Holyoke material, a new reconstruction is proposed that presents continentalTreptichnusas a shallow mole-tunnel-like burrow produced just below the sediment surface. This reconstruction is consistent with the morphology of RecentTreptichnus-like burrows produced by fly (dipteran) larvae, which are considered the most likely makers of the HolyokeTreptichnus.

Cryptic diversity in the Hypsiboas semilineatus species group (Amphibia, Anura) with the description of a new species from the eastern Guiana Shield.

We used molecular and morphological data to investigate the hidden diversity within the Hypsiboas semilineatus species group, and more specifically within H. geographicus, an allegedly widespread species in northern South America. As a result, the identity of H. geographicus was clarified, several candidate species were detected and one of them, from the eastern Guiana Shield, is described herein as a preliminary step to resolve the taxonomy of the group. Hypsiboas diabolicus sp. nov. is mainly distinguished from closely-related species by an acuminate snout in lateral view, well-developed webbing between fingers and toes, and unspotted carmine/crimson colouration on the concealed surfaces of legs, feet and hands in life. The tadpole of the new species is described and is characterized by a large A-2 gap, a mostly single row of large marginal papillae, and a dark brown to black colouration. We also describe the advertisement call of the new species, which is defined as a soft call consisting of short clusters of 2-3 chuckles with a dominant frequency ranging between 1.11-1.19 kHz. Hypsiboas diabolicus sp. nov. is currently known only from the eastern Guiana Shield, and is probably endemic to that region. The new species' range overlaps broadly with another candidate species referred to as H. aff. semilineatus 1. Our preliminary results stress out a high cryptic diversity in that species group and the need for a formal redescription of Hypsiboas geographicus based on more topotypic material than what is currently available to properly sort out the taxonomic status of several lineages in that clade.

Revision of the enigmatic Southeast Asian spider genus Savarna (Araneae, Pholcidae)

The genus Savarna Huber, 2005 was previously one of the most poorly known Pholcinae genera. Less than 20 specimens (representing four nominal species) were available worldwide; nothing was known about ultrastructure, natural history, or relationships. We present the first SEM data, supporting the position of the genus in Pholcinae outside the Pholcus group of genera and weakly suggesting a closer relationship with the genera Khorata Huber, 2005, Spermophorides Wunderlich, 1992, and two undescribed species of unknown affinity from Borneo. We provide the first data about microhabitat, web structure, and reaction to disturbance. We clarify the type locality of Savarna tessellata (Simon, 1901) (“Jalor, Biserat”) and describe topotypical material. We describe the previously unknown male of Spermophora miser Bristowe, 1952 and transfer the species (that was previously considered incertae sedis) to Savarna as Savarna miser (Bristowe, 1952) comb. nov. Savarna baso (Roewer, 1963) is newly synonymized with S. miser. We newly describe the most northern species in the genus, Savarna kaeo sp. nov., and provide amendments to the descriptions of all previously described species.

New fossiliferous sites with Barremian Charophyta in the "Grès du Liban" auct. (Lebanon), with a critical perspective regarding the nature of Munieria Deecke, 1883

The Gres du Liban auct. is the thick basal unit of the Lower Cretaceous series in Lebanon. There are two key levels in this dominantly siliciclastic unit: the Banc de Mrejatt and the Calcaire a pisolithes, the latter of which includes limestones and marls with Charophyta. The rich charophyte assemblage described by Grambast and Lorch (1968) from a site near Jezzine is Barremian (possibly Early Barremian) in age, not Bedoulian as previously stated. Besides gyrogonites and utricles, charophyte thalli are rather common; they are ascribed to two organ genera, Munieria Deecke, 1883, and Charaxis Harris, 1939. Based on topotypic material (from the Pia Collection), Munieria baconica Deecke, 1883, the type-species of the genus, is reassessed. In addition, the species Clypeina parvula Carozzi, 1946, is transferred to the revised genus Munieria, and a new species of Charaxis is described.

Two ammonite species under the same name: Revision of Deshayesites deshayesi (d’Orbigny, 1841) based on topotype material (Lower Aptian, Lower Cretaceous, Northeast of France)

After a review of the current state of knowledge about the classical Lower Aptian (Lower Cretaceous) ammonite index-species Deshayesites deshayesi (d’Orbigny, 1841) and the problems concerning the acceptation of this species, a topotype sample of 159 Deshayesites from the Argiles à Plicatules Formation (Paris Basin, northeastern France), including the lectotype of the latter species, is studied statistically. According to the Deshayesitidae evolutionary pattern highlighted by Bersac and Bert (2012), the present study shows that the whole studied population from the Argiles à Plicatules Formation belongs actually to a single species with a large intraspecific variability under the Dipolar Variation. Nothing supports the hypothesis of polyzonality of the level that provides the Deshayesites within the Argiles à Plicatules. The characteristics of the Deshayesites studied relate the latter to Deshayesites annelidus Casey, 1963 rather than to what is usually considered to be D. deshayesi. Consequently, D. deshayesi is revised, D. annelidus is treated as a junior synonym of D. deshayesi, and the species previously usually considered as D. deshayesi has to be renamed in D. multicostatus Swinnerton, 1935. Accordingly, it is proposed to move the limit of the D. deshayesi Zone downward.

Callichirus garthi is a valid species, distinct from C. seilacheri (Decapoda: Axiidea: Callianassidae)

Callichirus garthi from the southeastern Pacific has long been considered as a junior synonym of Callichirus seilacheri from the eastern tropical Pacific. The present study is based on a detailed comparison between both taxa collected along the coast of Chile and the Pacific coast of Central America, including type material of C. garthi and topotypic material of C. garthi and C. seilacheri. This reveals that C. garthi is a valid species, with a suite of morphological features that allow it to be separated from C. seilacheri. Callichirus garthi can primarily be distinguished from C. seilacheri by closely set eyestalks; the proximal margin of the terminal antennular segment exceeding the proximal margin of the last antennal segment with the dorsal margins of both the basal and intermediate antennular segment setose; the merus of the larger cheliped armed with a rough semicircular lobe over the anterior ventral margin and the ischium curved downwards without ornamentation on the ventral margin. On the basis of the present study, we therefore resurrect C. garthi as a valid species for the Chilean and Peruvian populations.http://zoobank.org/urn:lsid:zoobank.org:pub:6FB68C22-5C87-4331-8FCA-CC8994034F3C

Trilobodrilus itoi sp. nov., with a Re-Description of T. nipponicus (Annelida: Dinophilidae) and a Molecular Phylogeny of the Genus.

The marine interstitial annelid Trilobodrilus itoi sp. nov., the sixth member of the genus, is described on the basis of specimens collected intertidally at Ishikari Beach, Hokkaido, Japan; this is the second species in the genus described from the Pacific Rim. In addition, T. nipponicus Uchida and Okuda, 1943 is re-described based on fresh topotypic material from Akkeshi, Hokkaido, Japan. From both species, we determined sequences of the nuclear 18S and 28S rRNA genes, and the mitochondrial cytochrome c oxidase subunit I (COI) gene. Molecular phylogenetic trees based on concatenated sequences of the three genes showed that T. itoi and T. nipponicus form a clade, which was the sister group to a clade containing the two European congeners T. axi Westheide, 1967 and T. heideri Remane, 1925. The Kimura two-parameter distance for COI was 22.5-22.7% between T. itoi and T. nipponicus, comparable with interspecific values in other polychaete genera. We assessed the taxonomic utility of epidermal inclusions and found that the known six species can be classified into three groups.

Notes on Enidae, 6: On Euchondrus pseudovularis, an endemic species in Israel (Gastropoda, Pulmonata: Enidae)

A revision is given of a peculiar land snail species from Israel, previously classified as two different species known under the name Jaminia ovularis or Euchondrus aff. ovularis, and E. peudovularis. The species is mainly found on the kurkar ridges of the Coastal Plain, and is also known from a few scattered records from the northern part of the Negev Desert. The species turned out to be not identical with Multidentula ovularis, but a representative of the genus Euchondrus. A revision of the types and topotypical material of the recently described E. pseudovularis from the Negev revealed that the Israelian species known as Jaminia ovularis or Euchondrus aff. ovularis should be classified under E. pseudovularis. An overview is given of the history, ecology, taxonomy and distribution of this enid species endemic to Israel, and a list is given of the taxa that are currently classified within the genera Multidentula and Euchondrus.

Himalayan Cambrian brachiopods

AbstractA synoptic analysis of previously published material and new finds reveals that Himalayan Cambrian brachiopods can be referred to 18 genera, of which 17 are considered herein. These contain 20 taxa assigned to species, of which five are new: Eohadrotreta haydeni, Aphelotreta khemangarensis, Hadrotreta timchristiorum, Prototreta? sumnaensis and Amictocracens? brocki. Well‐preserved topotype material from the classic Parahio Valley section is described for three species that have not previously been illustrated photographically. A biostratigraphical scheme based on Cambrian brachiopod occurrence is integrated with that recently proposed for trilobites. Regional correlations based on shared taxa within and across Himalayan lithotectonic belts demonstrate that erosion of Cambrian rocks in the Kurgiakh orogeny in the Parahio Valley removed little of the Parahio Formation and that all of the fossil‐bearing lithotectonic zones share similar late early Cambrian brachiopods, suggesting regional faunal continuity at the time. Palaeobiogeographical analysis of the Cambrian brachiopod faunas from the Himalaya shows that they occupied a marginal position of the Australasian segment of Gondwana.

Redescription of Bythotrephes longimanus Leydig, 1860 and B. cederströmii Schödler, 1877 (Crustacea: Cladocera: Onychopoda), with notes on the morphology and systematics of the genus Bythotrephes Leydig, 1860.

Two core species of the genus Bythotrephes Leydig, B. longimanus Leydig and B. cederströmii Schӧdler, are redescribed on the basis of the type material, topotypic material, and other materials from Western and Northern Europe. They were investigated with an application of detailed morphological analysis, including intra- and interpopulation variability and the original scheme of morphometric measurements. The neotype of B. longimanus was selected from the topotypic population of the species from Bodensee (Switzerland-Germany). The taxonomy of B. cederströmii and of the whole genus is complicated by the presence of a variety of similar forms with intermediate morphology supposed to be interspecific hybrids, which much confused previous researches. The evolutionary transformations of some morphological structures, such as thoracic limbs, abdomen, and caudal process, are analyzed and discussed. In particular, it is suggested that the internal side of the endopodites of the thoracic limbs of the second-fourth pairs grew in proximal direction, occupying the part of protopodite with the "gnathobasic" process, which remains to be homologous to those of the thoracic limbs of the first pair. The strongly reduced thoracic limbs of the fourth pair lost two terminal segments and have proved to be represented by three segments, two proximal of which are those of protopodite, while the third terminal segment is composed of the fused distal part of the protopodite and the first proximal endopodital segment, bearing transformed and specifically arranged setae. The specificity of the "gnathobasic" processes of Bythotrephes is that they are the derivatives of the second, not of the first, inner endite of the thoracic limb and for this reason they are not homologues to the gnathobases of other cladocerans and should be called pseudognathobases.The same position of the "gnathobasic" processes is characteristic for other Onychopoda, which stresses the uniqueness of the representatives of the order among Cladocera and the whole class Branchiopoda. The abdomen of Bythotrephes was found to be three-segmented, but the third distal segment has proved to be transformed and fused with the postabdomen, participating in the formation of a long and massive caudal process. The comparison of morphological features of Bythotrephes and Leptodora clearly shows their independent homoplasic evolutionary development and distant phylogenetic position. According to new available data, the range of B. longimanus embraces large and deep lakes of Switzerland, Southern Germany, North Italy, Austria, and Great Britain, whereas B. cederströmiis. str. is certainly known only in the south of Sweden (probably also in Finland). All other B. cederströmii-like forms, supposed to be interspecific hybrids, are much more widely distributed in Eurasia from Northern Europe and Central European Russia to the Lower Volga River and Yakutia in Eastern Siberia. It is suggested that just these hybrids invaded and conquered the North American inland waters.

Review of the large branchiopod crustacean fauna of the Indian subcontinent (Anostraca, Notostraca, Laevicaudata, Spinicaudata, Cyclestherida)

The large branchiopod crustaceans (fairy shrimp, tadpole shrimp, and clam shrimp) of India have been examined with varying degrees of scientific rigour since the late 1800s. Although 86 nominal species have been reported for India, the validity of many is in question – particularly for the clam shrimp. Most Indian endemic taxa are poorly described; some lack type material or the type material has been lost. Most recent revisions of key groups have made no attempt to validate identifications using existing type material, or even to procure topotype material. Worse, recent reviews treat old synonyms as valid names without any analysis or explanation. We provide a review of the taxa from the Indian subcontinent (Bangladesh, India, Nepal, Pakistan, and Sri Lanka) taking a strongly conservative approach to assessing the reported species. We recognize only 42 species: 16 anostracans, 2 notostracans, 3 laevicaudatans, 21 spinicaudatans (with reservations), and a single species of cyclestherid. This should establish a base point for future work on the branchiopods of the subcontinent.

Redescription of Alitta succinea (Leuckart, 1847) and reinstatement of A. acutifolia (Ehlers, 1901) n. comb. based upon morphological and molecular data (Polychaeta: Nereididae).

The nereidid worm Alitta succinea (Leuckart, 1847), described from Western Germany, has been considered by some authors as a widespread and alien invasive species, or else as a group of morphologically indistinguishable species. Neither idea has yet been supported by critical taxonomic revisions of relevant material. Most characterizations of A. succinea were based upon a mixture of morphological features from specimens from the type locality and from other regions. Moreover, four species described from America are considered junior synonyms of A. succinea, including Nereis acutifolia Ehlers, 1901, described from the eastern tropical Pacific. The type material of the latter species has not been reviewed since its description. We re-examined type and topotype materials of A. succinea and N. acutifolia including atokous and epitokous specimens. In addition, newly collected specimens were used to evaluate genetic divergence between both species using the mitochondrial gene COI. Alitta succinea is redescribed from type material and specimens from Germany. We rejected the recent placement of the species in Neanthes and we transferred it to Alitta. Further, we refuse the synonymy of N. acutifolia with A. succinea due to morphological and molecular differences. Consequently, we regard Alitta acutifolia n. comb. as a valid species, and the supposed introduction and the alien status of A. succinea along the Mexican and Central American Pacific shores are rejected.

Intraspecific variation and taphonomic alteration in the Cambrian (Furongian) agnostoid Lotagnostus americanus: new information from China

The concept of the agnostoid arthropod species Lotagnostus americanus (Billings, 1860), which has been reported from numerous localities in the upper Furongian Series (Cambrian) of Laurentia, Gondwana, Baltica, Avalonia, and Siberia, is reviewed with emphasis on morphologic and taphonomic information afforded by large collections from Hunan in South China, Xinjiang in Northwest China, and Zhejiang in Southeast China. Comparisons are made with type and topotype material from Quebec, Canada, as well as material from elsewhere in Canada, the USA, the United Kingdom, Sweden, Russia, and Kazakhstan. The new information clarifies the limits of morphologic variability in L. americanus owing to ontogenetic changes and variation within holaspides, including inferred microevolution. It also provides details on apparent variation of taphonomic origin. The Chinese collections demonstrate a moderately wide variation in L. americanus, indicating that arguments favoring restriction of Lotagnostus species to narrowly defined, geographically restricted forms are unwarranted. Species described as L. trisectus (Salter, 1864), L. asiaticus Troedsson, 1937, and L. punctatus Lu, 1964, for example, fall within the range of variation observed in L. americanus, and are regarded as junior synonyms. The effaced form Lotagnosilis obscurus Palmer, 1955 is removed from synonymy with L. americanus. A review of the stratigraphic distribution of L. americanus as construed here shows that the earliest occurrences of the species in all regions of the world are nearly synchronous. (Less)

Neogoniolithon (Corallinales, Rhodophyta) on the Atlantic coast of Mexico, including N. siankanensis sp. nov.

This is the first revision of Neogoniolithon species on the east coast of Mexico based on both DNA sequences (psbA) and morpho-anatomical characters. Nine species names are provisionally applied based on morpho-anatomical comparisons with type specimens or with species descriptions: N. accretum, N. acropetum, N. erosum, N. mamillare, N. propinquum, N. rhizophorae, N. solubile, N. spectabile and N. strictum. For none of these species was type or topotype material sequenced, but all have their type localities in the subtropical or tropical western Atlantic Ocean. Neogoniolithon siankanensis is described on the basis of a distinctive branched thallus, apical tetrasporangial conceptacles and DNA sequence. Another four species are present, each characterized by DNA sequence, but insufficient reproductive material was available to adequately describe these species. A suite of vegetative and reproductive characters can adequately segregate the currently named Mexican east coast Neogoniolithon species, including 1) branched versus unbranched thalli, 2) monomerous versus dimerous construction, 3) disposition of trichocytes, 4) interior dimensions of mature tetrasporangial conceptacles, 5) number of cell layers comprising tetrasporangial conceptacle roof and 6) shape of cells that line tetrasporangial pore canal. Morpho-anatomical descriptions and distributions are provided for all named species.

Phylogenetic analysis of rhodolith formation in the Corallinales (Rhodophyta)

Although the ecological importance of rhodolith (maerl, free-living coralline algae) beds is well-known, rhodolith-forming species have been neglected in molecular phylogenetic studies. This is the first molecular systematic study aimed at understanding whether the rhodolith habit is a fixed feature in lineages and determining the relationship (phylogenetic vs. environmental) between rhodolith and crustose habits. Phylogenetic relationships of rhodolith-forming species and encrusting coralline algae at generic and species levels were analysed using SSU rDNA and psbA sequences. Extensive sampling in the European North Atlantic, Pacific and Caribbean Mexico of Phymatolithon, Lithothamnion, Lithophyllum and Neogoniolithon taxa forming rhodoliths and crusts was accompanied by examination of type or topotype material. Phylogenetic reconstruction showed that Neogoniolithon contained a monophyletic group of rhodolith-forming species whereas other rhodolith-formers were closely related to encrusting forms in the genera Phymatolithon, Lithothamnion, Mesophyllum, Hydrolithon, Spongites and Sporolithon. DNA analysis showed that the crust-forming Lithophyllum cf. incrustans/dentatum also forms rhodoliths with a stone nucleus that occur on rocky shores. In contrast, species that form beds of non-nucleate rhodoliths (e.g. Neogoniolithon spectabile, N. strictum, Lithophyllum cf. incrustans/dentatum or sp. 1 and Phymatolithon calcareum) rarely form crusts. The rhodolith habit cannot be used to delimit species for taxonomic or identification purposes. Extensive taxonomic revision will be required to deal with problems such as the position of specimens identified as Lithophyllum margaritae in two unrelated lineages.

The genus Brachyseris Alloiteau 1946/47 and remarks on Latimaeandraraea felixi Angelis d’Ossat, 1905 (Scleractinia; Cretaceous)

The Early Albian coral species Latimaeandraraea felixi is investigated on the basis of thin sections obtained from the holotype. Its fine skeletal structure and systematic position are elucidated, but its generic position remains questionable. It is most closely related to the genus Brachyseris from which it differs by lacking the typical pattern (forming tholiform walls in two different levels) of this genus. The genus Brachyserisis considered in greater detail on the basis of the holotype and topotypical material of the type species. It is related to other Leptophyllid genera such as Microphyllia, Thalamocaeniopsis, Trigerastraea and Vallimeandra, but is very probably restricted to the Late Cretaceous.

Studies on species of Holarctic Pardosa groups (Araneae, Lycosidae). VIII. The Palearctic species of the Pardosa nigra group.

The eight Palearctic species of the Pardosa nigra group are reviewed, illustrated and keyed. Two of them are redescribed: Pardosa lyrata (Odenwall, 1901) and P. paramushirensis (Nakatsudi, 1937). The former is widespread in Siberia and has also been found in Mongolia while the latter has been recorded from Japan (Honshu and Hokkaido) and Russia: the Kuril Islands (a neotype is designated from topotypic material from Paramushir Island). Pardosa fomichevi sp. n. is described from the Maritime Province (Primorye) in Russia, based on both sexes. Pardosa arctica (Kulczyński, 1916) is placed as a synonym to P. eiseni (Thorell, 1875).