Research Article |
Corresponding author: Andreas Schmidt-Rhaesa ( a.schmidt-rhaesa@leibniz-lib.de ) Academic editor: Michael Ohl
© 2016 Andreas Schmidt-Rhaesa.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Schmidt-Rhaesa A (2016) The collection of Nematomorpha in the Zoological Museum Hamburg, including description of a new species, Chordodes jelkae sp. n. Zoosystematics and Evolution 92(2): 211-250. https://doi.org/10.3897/zse.92.10351
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The collection of horsehair worms in the Zoological Museum of the Centrum für Naturkunde (CeNak) of the University of Hamburg has been revised. All specimens have been investigated by Scanning Electron Microscopy (SEM). A total of 173 specimens from 135 catalogue entries is present in the collection, these represent 39 species from 10 genera (Gordius, Chordodes, Paragordius, Parachordodes, Gordionus, Acutogordius, Beatogordius, Paragordionus, Pseudochordodes, Nectonema). Previous revisions of the Nematomorpha material have been done in 1893 by Römer and in 1935 by Heinze. A number of species could not be determined to species level, either because characters were not preserved well enough or because observed characters did not fit available species descriptions. This does account in particular for the genus Gordius, where diagnostic characters are few and great uncertainties exist concerning the status of certain species. Therefore, within this genus, many records were only determined as Gordius sp., including some specimens that had previously been determined to species level. One new species is described from the material of the collection. This species is named Chordodes jelkae sp. n. and it is characterized by the presence of an apical tuft of bristles on simple areoles in combination with the absence of thorn areoles. The holotype of Chordodes pilosus Möbius, 1855 was reinvestigated, and its species identity cannot be determined unambiguously. Therefore the name Chordodes pilosus is considered a nomen dubium.
Nematomorpha , Chordodes jelkae sp. n., horsehair worms, collection, revision, SEM
The Zoological Museum Hamburg has a collection of 173 specimens from 135 catalogue entries of horsehair worms (Nematomorpha) representing 39 species from 10 genera. A total of about 455 species has been described from all over the world. Important for collection and determination are adult worms in their free-living phase (see
The only published review of the collection of the Zoological Museum Hamburg is from
In 1935, Kurt Heinze from Berlin worked on the Hamburg collection and determined a number of specimens, a few of these results were published in
Since then, nobody has reviewed the material or deposited new type material until from 2008 on the author deposited several types in the collection.
It is the aim of this summary on the one hand to present an overview of the Nematomorpha material in the Hamburg collection and on the other hand to determine all material according to current standards. This includes investigation by Scanning Electron Microscopy (SEM). The SEM reinvestigation led to several new determinations, in particular a number of determinations at the species level could not be confirmed and hence are treated here as “sp.”.
Specimens are conserved in ethanol with a desired concentration of 70%. For the reinvestigation, entire pieces from the midbody region and in some cases the posterior end (about 1–2 mm from the terminal tip) were prepared for Scanning Electron Microscopy (SEM). Pieces were dehydrated in an increasing ethanol series, critically point dried and coated with gold in a sputter coater. Observation took place using a LEO SEM 1524 at 10 kV. Digital images were taken.
In the Zoological Museum Hamburg, entries are recorded in a catalogue book, printed as a record card and recorded digitally in a Filemaker file. Entries in the catalogue are hand-written and could not always be read reliably. It was attempted to verify the collection localities, but this was not possible in all cases. Writing of names is given here as good as possible. Citations from the catalogue are given in quotation marks. The main focus is on the older records, which are reinvestigated. Younger records, which are well documented in the literature, are presented only briefly, with reference to the respective publications.
Males in species of Acutogordius have a postcloacal crescent, like in Gordius, but the tail lobes are pointed and the entire posterior end tapers towards the pointed tip of the tail lobes (Fig.
Acutogordius cf. acuminatus De Miralles & De Villalobos, 1998
Material examined. V 2213 (1 male). Fig.
Location. Brazil, Joinville, Santa Catarina.
Remarks from the catalogue. Collected by "Killmann" (no date given). Received by the museum on June 19, 1890. Determined by Römer in 1893 as Gordius aquaticus and by Heinze in 1935 as Gordius sp.
Occurs in publications.
Remarks.
Material examined. V7351(b) (1 male). Fig.
Location. Brazil, presumably Itapoá, Santa Catarina (see remarks from the catalogue).
Remarks from the catalogue. Collected by W. Ehrhardt on January 17, 1910. Received by the museum on November 12, 1910. As location, the catalogue says “Humboldt-Geb. am Itapoeú", some letters are not explicit. This location could not be retrieved, but Itapoá is close. Additionally, there are other catalogue entries with the same date and same collector having as location Joinville, which is close to Itapoá in the province Santa Catarina. Original determination as "Gordiide".
Occurs in publications. none
Remarks. The catalogue notes “dv”, meaning “diverse” as number of specimens. 13 specimens were found, 12 of them are determined as Chordodes brasiliensis (see there) and specimen V7351(b) belongs to the genus Acutogordius. No bristles were observed on the body cuticle (Fig.
Material examined. V11465 (1 male). Fig.
Location. Brazil, Boiteuxburgo, Santa Catarina.
Remarks from the catalogue. Collected by Paul Missfeldt (date unknown). Received by the museum on December 5, 1930. Originally determined as “Gordius”.
Occurs in publications. unknown
Remarks. The specimen has clearly pointed tail lobes, which carry two longitudinal crests or keels (Fig.
Characteristic for this genus, which currently includes 21 species (
Beatogordius sp.
Material examined. V8939 (1 male). Fig.
Location. Chile, Coronel, in drinking water.
Remarks from the catalogue. collected by R. Paeseler on October 5, 1914. Received by the museum on January 15, 1920. Originally determined as “Gordiide”.
Occurs in publications. unknown
Remarks. The cuticle is not very well preserved and strongly covered with dirt. Paired elongate areoles arranged in longitudinal rows can be well observed (Fig.
Material examined. V9575 (1 male). Fig.
Location. Chile, Limache (writing is not explicit), in river.
Remarks from the catalogue. collected by D.C. Bock (date not given). Received by the museum on July 9, 1923. Originally determined as “Gordiide”
Occurs in publications. unknown
Remarks. The cuticle of this specimen shows longitudinal rows of paired, elongate areoles, which is characteristic for the genus Beatogordius (Fig.
Beatogordius chinensis Schmidt-Rhaesa, 2011
Material examined. V13294 (1 female)
Location. China, Yunnan province, 2 km south of Haba, Haba Xueshan Mountains (27°21.9'N, 100°08.3'E), 2870m. Collected in a small pond in a dry valley of a brook.
Remarks from the catalogue. Collected by J. Hájek and J. Růžička on June 17–20, 2007.
Occurs in publications.
Remarks. Holotype and only known specimen as described in
This is a genus with almost 100 described species, most of which occur in the tropics and subtropics and many of which have mantids as host (see
Chordodes sp.
Material examined. V2230 (1 female). Fig.
Location. Venezuela, Ciudad Bolivar.
Remarks from the catalogue. Collected by D. Siegert (no collection date or date when received by the museum given). Determined by Möbius in 1855 as Chordodes pilosus, confirmed by Römer in 1893 and Heinze in 1935. Host: Blabera gigantea.
Occurs in publications.
Remarks. This specimen is the holotype for the species Chordodes pilosus, described by Möbius in 1855. The genus Chordodes had been introduced briefly before (
Material examined. V5320 (1 male, 1 female). Fig.
Location. South Africa, Bothaville.
Remarks from the catalogue. Collected by Brauns (no date given). Received by the museum on November 13, 1899. Determined as “Gordius”. From “Mantis”.
Occurs in publications. unknown
Remarks. The cuticle of the female is in comparably bad shape. Crowned areoles with short and long apical filaments are present (Fig.
Chordodes brasiliensis Janda, 1893
Material examined. V7351 (a) and (c-m) (10 males and 2 females). Fig.
Location. Brazil, presumably Itapoá, Santa Catarina (see remarks from the catalogue).
Remarks from the catalogue. Collected by W. Ehrhardt on January 17, 1910. As location, the catalogue says “Humboldt-Geb. am Itapoeú", some letters are not clear. This location could not be retrieved, but Itapoá is close. Additionally, there are other catalogue entries with the same date and same collector having as location Joinville, which is close to Itapoá in the province Santa Catarina.
Occurs in publications. None
Remarks. The catalogue notes “dv”, meaning “diverse” as number of specimens. Thirteen specimens were found, one of them (b) is determined as Acutogordius cf. acuminatus (see there), the others are determined as Chordodes brasiliensis. The cuticle has numerous isolated, tall and slender crowned areoles (Fig.
Chordodes jandae Camerano, 1895 or C. skorikovi Camerano, 1903
Material examined. V4064 (1 male and 1 female). Fig.
Location. Indonesia, Sumatra (as “O. Sumatra, Dali, Indrapura Estab”).
Remarks from the catalogue. Collected by “Gebr. Siemssea” (no date given). Received by the museum on March 21, 1896. Determined as “Gordiiden”.
Occurs in publications. unknown
Remarks. The simple areoles have the shape of blackberry areoles (Fig.
Chordodes janovyi Bolek, Schmidt-Rhaesa, Hanelt & Richardson, 2010
Material examined. V13291 (1 male)
Location. Cameroon, Menoua river drainage in the village Bawa (5°24'N, 10°03'E).
Remarks from the catalogue. Collected in June 2006.
Occurs in publications.
Remarks. Holotype of this species. See
Material examined. V13292
Same data as for V13291, paratype of this species. See
Chordodes japonensis Inoue, 1951
Material examined. V4599 (sex unknown because of damaged posterior end). Fig.
Location. “Japan, Schikoku, Tosa, Mano Bogawa, Kamigori”
Remarks from the catalogue. Collected by Lenz on July 27, 1897. Received by the museum on August 11, 1897. Determined as “Gordiide”.
Occurs in publications. unknown
Remarks. The cuticle of the specimen is in most of the investigated part covered by the unstructured larval cuticle (Fig.
Material examined. V13282 (female).
Location. Japan, Campus of Nakamura Gakuen Sanyo High School, Imajukuaoki 1042-33, Nishi-ku, Fukuoka City, Fukuoka Prefecture, Kyushu (33°33'58"N, 130°17'22"E).
Remarks from the catalogue. Collected by Yuriko Kuribashi on October 12-18, 2008 from host Hierodula patellifera.
Occurs in publications.
Remarks. This specimen is only briefly mentioned in
Chordodes cf. japonensis
Material examined. V5202 (1 male). Fig.
Location. Japan, Tokyo
Remarks from the catalogue. Collected by Jourdan (no date given). Received by the museum on February 14, 1899. Found in an undetermined mantid. Determined as “Gordius”.
Occurs in publications. unknown
Remarks. The cuticle is not in very good shape, but it largely corresponds to the cuticular pattern of C. japonensis (compare specimen V4599 and
Chordodes cf. joyeuxi Dorier, 1935
Material examined. V5778 (1 female). Fig.
Location. Indonesia, Sumatra, Lomgei Lalah, Indragiri
Remarks from the catalogue. Collected by W. Burchard (no date given). Received by the museum on August 26, 1901. Determined as “Gordius”.
Occurs in publications. unknown
Remarks. The simple areoles have the shape of blackberry areoles (Fig.
Chordodes parabipilus Kintsurashvili, Schmidt-Rhaesa & Gorgadze, 2011
Material examined. V13269 (1 female, holotype)
Location. Georgia, village Tsitamuri, 20 km from Tbilisi.
Remarks from the catalogue. Collected by Temur Gvalia on September 30, 2002, given by N. Kintsurashvili. Determined by Schmidt-Rhaesa in 2008.
Occurs in publications.
Remarks. Holotype of this species. See
Chordodes polycoronatus Schmidt-Rhaesa & Brune, 2008
Material examined. V11540 (1 male). Fig.
Location. Langcat, Sumatra, Indonesia.
Remarks from the catalogue. Collected by Ernst Versmann (date unknown). Received by the museum on March 9, 1931. Originally determined as “Gordiide”.
Occurs in publications. unknown
Remarks. With the large clusters of crowned areoles (Fig.
Chordodes queenslandi Schmidt-Rhaesa, 2002
Material examined. V2231 (1 female)
Location. Australia, Sidney.
Remarks from the catalogue. Collected by Dämel (no collection date or date when received by the museum given). Determined by Römer in 1893 as C. bouvieri, by Heinze in 1935 as C. modiglianii and by Schmidt-Rhaesa in 2002 as C. queenslandi.
Occurs in publications.
Remarks. As explained in
V10960 (2 females). Fig.
Rwanda, Kabgayi, “Poste de Kigali”.
Specimen 10960 (I).
Collected by "Freres Rodrigues v.d. Weissen Vätern" (no collection date). Received by the museum on September 20, 1929.
The species is named after and dedicated to my oldest daughter, Jelka.
The two females measure 90 (I) and 85 (II) mm in length and 0.9/1.0 mm in diameter, respectively. The anterior end tapers towards the tip. The body colour is a dark brown general occurrence, with brown basic colour and darker patches (the “leopard pattern”). The anterior tip is white and blends into the brown body colour. Specimen (I) is well preserved, characters of specimen (II) are less well visible (Fig.
Species of Chordodes are quite similar in their cuticular structure. Species are distinguished by the presence or absence of particular types of areoles or when cuticular structures exhibit a particular substructure. Characteristic for this new species is the form of the simple areoles. In most species of Chordodes the simple areoles have either no bristles or scattered fine bristles on their apical surface. SEM investigations reveal that small bristles may be more abundant than known on the basis of traditional light microscopical investigation and they may even form small tuft-like structures as present, e.g. in C. parabipilus Kintsurashvili, Schmidt-Rhaesa & Gorgadze, 2011 (
The genus Gordionus is characterized by the following character combination: one type of areoles present on the cuticle, male posterior end with bristles anterolateral of the cloacal opening and spines posterior of the cloacal opening and directly around it. Currently, 56 species are included in this genus (
Gordionus sp.
Material examined. V2215 (3 females and one specimen without posterior end). Fig.
Location. “East Africa, middle Ussuri” (as “Ostafrika, mittlerer Ussuri”) [a river Ussuri could not be located].
Remarks from the catalogue. Collected by “Fischer” (no collection date or date when received by the museum given). Carabus smaragdinus is given as host. Determined by Römer in 1895 and Heinze in 1935 as male of Gordius aquaticus.
Occurs in publications.
Remarks. Presently, the vial contains four specimens: three females, of which one was investigated here by SEM and one specimen in an extra vial, this specimen is lacking the posterior end. According to the catalogue the number of specimens should be 2 and the determination says “Gordius aquaticus ♂”.
Material examined. V2217 (according to catalogue 3 males, only 1 male found in vial). Fig.
Location. Germany (not further specified), from carabid beetle.
Remarks from the catalogue. Collected by “v. Siebold” (no collection date or date when received by the museum given). Determined by Von Siebold as Gordius subbifurcatus, by Römer in 1893 as Gordius tolosanus. Genus corrected to Parachordodes by Heinze in 1935 (according to additional label in vial). Additional remark: type material of Gordius subbifurcatus.
Occurs in publications.
Remarks. Gordius subbifurcus Von Siebold, 1848 (not subbifurcatus as in catalogue!) is synonymous to Parachordodes tolosanus (Dujardin, 1842) (see, e.g.,
Finally, the determination as Parachordodes tolosanus could not be supported here. The genus Parachordodes has two types of areoles, one of which are the so-called superareoles (see, e.g.,
Material examined. V2236 (II) (1 female) One specimen is indicated in the catalogue, but 2 specimens are present. The second specimen (I) is determined as Parachordodes tolosanus. Fig.
Location. Germany, Göttingen.
Remarks from the catalogue. Collected by Schlotthauber (no collection date or date when received by the museum given). Determined by Schlotthauber in 1854 as Gordius aquaticus, corrected by Heinze in 1935 to Parachordodes tolosanus. Host: Harpalus ruficornis crossed out and replaced by Ophonus cephalotes.
Occurs in publications (as P. tolosanus).
Remarks. Only one type of areoles is present (Fig.
Material examined. V9615 (2 females) (3 specimens are indicated in the catalogue). Fig.
Location. Macedonia, stream to river Vardar, close to Veles (as “Bach zum Vardoe, N. v. Veles”)
Remarks from the catalogue. collected by F. Ahlborn in 1916 (date not given). Received by the museum on November 22, 1923. Originally determined as “Gordius”.
Occurs in publications. unknown
Remarks. The cuticle of both specimens has roundish areoles surrounded by a broad and flat interareolar space, in which interareolar bristles are present (Fig.
Material examined. V11117 (1 male). Fig.
Location. Germany, ditch in Lokstedt, Hamburg.
Remarks from the catalogue. Collected by H. Richter (date unknown). Received by the museum on July 25, 1930. Determined by Heinze in 1935 as Gordionus scaber.
Occurs in publications.
Remarks. The areoles on the cuticle of this specimen are not clearly recognizable and appear to be artificially altered (Fig.
Material examined. V12470 (1 male). Fig.
Location. Germany, Friedrichstadt an der Eider.
Remarks from the catalogue. collected by Niggemeyer (no date given). Received by the museum on May 16, 1936. Determined by Thiel in 1936 as Gordius aquaticus.
Occurs in publications. unknown
Remarks. The posterior end of this specimen is unfortunately lost during preparation for SEM, but notes taken before the loss indicate that a postcloacal crescent is absent. The cuticle has one type of areoles (Fig.
Material examined. V13313 (1 male)
Location. Japan, Hirakura, Higaashim.
Remarks from the catalogue. Collected by Takuya Sato on September 12, 2008. Received by the museum in 2011.
Occurs in publications.
Remarks. Specimen is a voucher for a phylogenetic analysis in
Material examined. V13316 (1 male)
Location. Japan, Hirakura, Nishimata.
Remarks from the catalogue. Collected by Takuya Sato on September 29, 2008. Received by the museum in 2011.
Occurs in publications.
Remarks. Specimen is a voucher for a phylogenetic analysis in
Material examined. V13319 (1 male)
Location. Japan, Kumano, Misen.
Remarks from the catalogue. Collected by Takuya Sato on September 14, 2009. Received by the museum in 2011.
Occurs in publications.
Remarks. Specimen is a voucher for a phylogenetic analysis in
Material examined. V13320 (1 male)
Location. Japan, Hirakura, Nishimata.
Remarks from the catalogue. Collected by Takuya Sato on September 14, 2009. Received by the museum in 2011.
Occurs in publications.
Remarks. Specimen is a voucher for a phylogenetic analysis in
Material examined. V13361 (1 female; transferred from V2220). Fig.
Location. Germany, Göttingen, Hainholz, in source.
Remarks from the catalogue. Collected by Schlotthauber (no collection date or date when received by the museum given). Determined by Römer in 1893 as Gordius violaceus, corrected by Heinze in 1935 to Parachordodes tolosanus.
Occurs in publications.
Remarks.SEM reinvestigation showed that the female does not have superareoles as required for the original determination as P. tolosanus. Because it has only one type of areoles (Fig.
Material examined. V13362 (1 male; transferred from V2225). Fig.
Location. Germany, Hamburg, river Bille.
Remarks from the catalogue. Collected by Steinblinck (no date given), received by the museum on January 1, 1895. Determined by Römer in 1893 as Gordius violaceus, corrected to Gordionus meissneri by Heinze in 1935.
Occurs in publications.
Remarks.SEM reinvestigation shows partly fused areoles (Fig.
Gordionus bageli Schmidt-Rhaesa & Gusich, 2010
Material examined. V13278 (1 male)
Location. Switzerland, Röserental near Liestal, Tafeljura. In Thuja forest.
Remarks from the catalogue. Collected by Valeria Viktoria Gusich in 2008.
Occurs in publications.
Remarks. Holotype for this species. See
Gordionus barbatus Schmidt-Rhaesa & Cieslak, 2008
Material examined. V13259 (1 male)
Location. Spain, Lleida, Llesp, Barranc Basculina (42°27'24.5"N, 0°44'57"W).
Remarks from the catalogue. Collected by J. Fresneda, I. Ribera and A. Cieslak on August 2, 2006. Received by the museum on July 3, 2008.
Occurs in publications.
Remarks. Holotype of this species. See
Gordionus chinensis (Villot, 1874)
Material examined. V13250 (1 male)
Location. Japan, Yumitehara river, Totsu river system, Kii Peninsula.
Remarks from the catalogue. Collected by Takuya Sato in 2007. Received by the museum on May 20, 2008.
Occurs in publications.
Remarks. see
Material examined. V13251 (1 male)
Same data as V13250, see there.
Material examined. V13312 (1 male)
Location. Japan, Arida, Ninomata.
Remarks from the catalogue. Collected by Takuya Sato on September 13, 2008. Received by the museum in 2011.
Occurs in publications.
Remarks. Specimen is a voucher for a phylogenetic analysis in
Material examined. V13314 (1 male)
Location. Japan, Hirakura, Higaashim.
Remarks from the catalogue. Collected by Takuya Sato on September 12, 2008. Received by the museum in 2011.
Occurs in publications.
Remarks. Specimen is a voucher for a phylogenetic analysis in
Material examined. V13315 (1 male)
Location. Japan, Miyagawa, Muzutani.
Remarks from the catalogue. Collected by Takuya Sato on September 29, 2008. Received by the museum in 2011.
Occurs in publications.
Remarks. Specimen is a voucher for a phylogenetic analysis in
Material examined. V13317 (1 male)
Location. Japan, Kumano, Sankou.
Remarks from the catalogue. Collected by Takuya Sato on October 13, 2010. Received by the museum in 2011.
Occurs in publications.
Remarks. Specimen is a voucher for a phylogenetic analysis in
Material examined. V13318 (1 male)
Location. Japan, Kumano, Sankou.
Remarks from the catalogue. Collected by Takuya Sato on October 13, 2010. Received by the museum in 2011.
Occurs in publications.
Remarks. Specimen is a voucher for a phylogenetic analysis in
Material examined. V13321 (1 male)
Location. Japan, Kumano, Sankou.
Remarks from the catalogue. Collected by Takuya Sato on October 13, 2010. Received by the museum in 2011.
Occurs in publications.
Remarks. Specimen is a voucher for a phylogenetic analysis in
Gordionus kii Schmidt-Rhaesa & Sato, 2009
Material examined. V13252 (1 male)
Location. Japan, Yumitehara river, Totsu river system, Kii Peninsula.
Remarks from the catalogue. Collected by Takuya Sato in 2007. Received by the museum on May 20, 2008.
Occurs in publications.
Remarks. Holotype for this species. See
Gordionus scaber Müller, 1926
Material examined. V2221 (1 male). Fig.
Location. Germany, Hamburg, Bergedorf.
Remarks from the catalogue. Collected by Thalenhorst (no collection date or date when received by the museum given). Determined by Römer in 1893 as Gordius violaceus and by Heinze in 1935 as Gordionus scaber (s. str.).
Occurs in publications.
Remarks. Although not optimally preserved, it is evident that the areoles are isolated or partly fused in short rows (Fig.
Material examined. V10924 (1 male). Fig.
Location. Germany, Medingen, Bad Bevensen, presumably stream Wohbeck (as “aus dem Wohbach”, “Wohbeck is the current name).
Remarks from the catalogue. Collected by F. Richter (date not given). Received by the museum on June 29, 1929. Determined by Heinze in 1935 as Gordionus scaber.
Occurs in publications.
Remarks. The cuticle shows roundish areoles with an individual character, but which are partly fused with neighbouring areoles in longitudinal direction (not figured). This is consistent with the pattern described for Gordionus scaber (e.g.,
Gordionus turkensis Schmidt-Rhaesa & Cieslak, 2008
Material examined. V13262 (1 male)
Location. Turkey, Karabük Rd. E80, E cross to Gerede, stream in Cayorenguney (40°48'23'N, 32°16'.5"E), 1231 m.
Remarks from the catalogue. Collected by I. Ribera on April 28, 2006. Received by the museum on July 3, 2008.
Occurs in publications.
Remarks. Holotype of this species. See
Gordionus violaceus (Baird, 1853)
Material examined. V2225 (originally 2 males, 1 male is regarded as Gordionus sp. and transferred to a new catalogue number 13362). Fig.
Location. Germany, Hamburg, river Bille.
Remarks from the catalogue. Collected by Steinblinck (no date given). Received by the museum on January 25, 1895. Determined by Römer in 1893 as Gordius violaceus, corrected to Gordionus meissneri by Heinze in 1935.
Occurs in publications.
Remarks.SEM reinvestigation shows one type of areoles surrounded by interareolar bristles (Fig.
Material examined. V11455 (2 females). Fig.
Location. Germany, Rügen.
Remarks from the catalogue. Collected by A. Thienemann (date unknown). Received by the museum on November 12, 1930. Originally determined by A. Thienemann as Parachordodes tolosanus. Corrected by Heinze in 1935 to Gordionus violaceus.
Occurs in publications. unknown.
Remarks. In both females the cuticle has round areoles completely surrounded by interareolar bristles (Fig.
Gordionus wolterstorffii (Camerano, 1888)
Material examined. V2218 (2 males, 1 female according to catalogue, presently only 1 male in vial). Fig.
Location. Germany, Göttingen, Hainholz, in source.
Remarks from the catalogue. Collected by Schlotthauber (no collection date or date when received by the museum given). Determined by Römer in 1893 as Gordius tolosanus, corrected to Gordionus scaber scaber by Heinze in 1935.
Occurs in publications.
Remarks. Collector and collection dates are similar to V2220 determined as Parachordodes tolosanus. The cuticle shows irregular rows of fused areoles (Fig.
Material examined. V2222 (in the catalogue 5 males and 2 females are indicated, presently only 1 male is present). Fig.
Location. Germany, Kissingen (additional remark "Poppenhausen" in
Remarks from the catalogue. Collected by Dömling (no date given). Received by the museum on September 23, 1893. Determined by Römer in 1893 as Gordius violaceus and by Heinze in 1935 as Gordionus scaber.
Occurs in publications.
Remarks. The cuticle shows irregular rows of fused areoles (Fig.
Material examined. V5255 (1 male). Fig.
Location. Germany, Hamburg, Elbe river.
Remarks from the catalogue. Collected by Wiebelitz (no date given). Received by the museum on June 14, 1899. Determined by Heinze in 1935 as Gordionus scaber.
Occurs in publications. unknown
Remarks. The cuticle shows irregular rows of fused areoles (Fig.
Species of the genus Gordius are not easy to determine due to their scarceness of diagnostic characters, especially in female worms. Additionally, the range of intraspecific variation for several characters is not known (
Characteristic for the genus is the presence of a semicircular or parabolic cuticular fold posterior of the ventral cloacal opening, which is called the postcloacal crescent. Posterior of this structure, the body is divided into paired lateral tail lobes. In Gordius, the tail lobes are rounded, in contrast to the genus Acutogordius, which has pointed tail lobes and is the only other genus with a postcloacal crescent (see below). In females, there is no difference to females of other genera, therefore females are more difficult to assign. The body cuticle in Gordius specimens can be either smooth or ornamented with roundish or polygonal structures called areoles. Areoles are present in other genera, too, especially in Gordionus. A smooth cuticle is only present in Gordius, but it has been described from several species (15 European species according to
Gordius sp.
Material examined. V 2207 (1 male). Fig.
Location. Germany, Kellersee, Holstein.
Remarks from the catalogue. Collected by "v. Pöppinghausen" (no date given), received by the museum on August 10, 1894. Determined as G. aquaticus by Römer in 1893 and as G. mülleri by Heinze in 1935.
Occurs in publications.
Remarks. In Gordius muelleri Heinze, 1933 (writing in catalogue mülleri) there is some distance between the postcloacal crescent and the beginning of the bifurcation of the tail lobes (see
Material examined. V 2208 (3 specimens, 1 male was reinvestigated here). Fig.
Location. Germany, Plön, Holstein.
Remarks from the catalogue. Collected by “Dr. G. Duncker” (no date given), received by the museum on January 18, 1894. Determined as G. aquaticus by Römer in 1893 and as G. albopunctatus by Heinze in 1935.
Occurs in publications.
Remarks. The cuticle is, in the investigated part, covered with material of unknown nature (Fig.
Material examined. V2214 (1 female). Fig.
Location. Chile
Remarks from the catalogue. No detailed collecting locality in Chile mentioned, collector unknown. No collection date given and no date when received by the museum. Determined by Römer in 1893 as G. aquaticus, confirmed by Heinze in 1935.
Occurs in publications.
Remarks. Female worm with smooth cuticle (Fig.
Material examined. V2216 (1 female). Fig.
Location. Chile, Valdivia river.
Remarks from the catalogue. In catalogue: “Coll. Mich. 33”, on record card “Michaelsen”. No further collection date or date when received by the museum given. Determined by Römer in 1893 as Gordius aquaticus, also mentioned “Heinze 1935”.
Occurs in publications.
Remarks. Female worm with smooth cuticle (Fig.
Material examined. V2223 (in the catalogue 1 male and 1 female are mentioned, currently two fragments are present, from which at least one is from the male). Fig.
Location. Germany, Hamburg (no further details).
Remarks from the catalogue. Collector unknown, no date given. Date when received by the museum not given. Determined by Römer in 1893 as Gordius violaceus and by Heinze in 1935 as Gordius mülleri.
Occurs in publications.
Remarks. The SEM reinvestigation shows a slight pattern of areoles on the cuticle (Fig.
Material examined. V2227 (1 female). Fig.
Location. “Viti Islands” (probably Viti Levu, Fiji-Islands).
Remarks from the catalogue. Collected or donated by “M.G.”. No collection date or date when received by the museum given. Host: Phybalosoma pythonis (Phasmatodea). Determined by Römer in 1893 as Gordius aeneus, determination checked and confirmed by Heinze in 1935.
Occurs in publications.
Remarks. Gordius aeneus is regarded an invalid species name (
Material examined. V2228 (1 female). Fig.
Location. unknown (“?” in catalogue).
Remarks from the catalogue. Collector unknown. No collection date or date when received by the museum given. Determined by Römer in 1893 as Gordius fulgur, determination confirmed by Heinze in 1935.
Occurs in publications.
Remarks.SEM investigation of specimen V2228 shows a smooth cuticular surface (Fig.
Material examined. V2238 (1 specimen of unknown sex). Fig.
Location. Tanzania, Bagamoyo.
Remarks from the catalogue. Collected by Stuhlmann. No collection date or date when received by the museum given. Determined first as Gordius aquaticus, but “aquaticus” was later crossed out. No name of reviser is given. Worm found in an undetermined cricket.
Occurs in publications. unknown
Remarks. The worm is still inside the cricket, some loops of it are visible through an injury in the anterior part of the abdomen. Body ends were not removed, and a sex determination is therefore not possible. A piece of worm was removed from the outermost loop. The cuticle is smooth (Fig.
Material examined. V2977 (1 female). Fig.
Location. Japan, Lake Candidius.
Remarks from the catalogue. Collected by H. Sauter (no date given), received by the museum on April 22, 1908. Originally determined as “Chordodes”.
Occurs in publications. unknown
Remarks.SEM investigation reveals a smooth cuticle (Fig.
Material examined. V3314 (1 female). Fig.
Location. “Elsass, Tannenberg bei Saales, 640m, in einer Heuschrecke”. Revised location: France, Saales. Tannenberg is probably a sanatorium close to Saales.
Remarks from the catalogue. Collected by Dr. Alb. Graser (no date given), received by the museum on July 12, 1909. Host: a cricket.
Occurs in publications. unknown.
Remarks. The cuticle is smooth, but contains small keel-like structures perpendicular to the longitudinal body axis (Fig.
Material examined. V4413 (1 male). Fig.
Location. Germany, “Bergedorf” close to Hamburg.
Remarks from the catalogue. Collected by “W. Fischer” (date not given). Received by the museum on October 1, 1896, determined as “Gordiide”.
Occurs in publications. unknown
Remarks. The cuticle contains shallow areoles with polygonal or irregular outline (Fig.
Material examined. V4598 (sex unknown, because posterior end is missing). Fig.
Location. Japan, Schikoku, Mano bogawa, Kamigore, Tosa.
Remarks from the catalogue. Collected by Lenz on July 27, 1897. Received by the museum on August 11, 1897. Originally determined as “Gordiide”.
Occurs in publications. unknown
Remarks. The specimen has a smooth cuticle (Fig.
Material examined. V4611 (1 male, 1 female). Fig.
Location. Italy, South Tyrol ("Süd-Tirol").
Remarks from the catalogue. Collected by Timm (no date given), received by the museum on August 21, 1897. Determined as “Gordius”.
Occurs in publications. unknown
Remarks. Both specimens differ in their cuticular structure, which may indicate that they belong to different species. The female has a smooth cuticle (not figured), while the cuticle of the male has elongate areoles that are partly confluent with neighbouring ones and that are arranged perpendicular to the longitudinal body axis (Fig.
Material examined. V4981 (1 male). Fig.
Location. Germany, Hamburg, Flottbeck, in drainage tubes.
Remarks from the catalogue. Collected by Ansorge (no date given), received by the museum on November 19, 1897. Determined by Heinze in 1935 as G. aquaticus.
Occurs in publications.
Remarks. The SEM reinvestigation shows a smooth cuticle (Fig.
Material examined. V5046 (1 female). Fig.
Location. Switzerland, “Axenstr. bei Tells Platte” (probably close to Sisikon, Uri, Swizerland).
Remarks from the catalogue. Collected by “Michaelsen 20.VI.98”, received by the museum on June 30, 1898. Determined as “Gordius”. Found in man (“im Menschen”).
Occurs in publications. unknown
Remarks. The cuticle is smooth, but has some fungus-like covering (Fig.
Material examined. V5459 (1 male). Fig.
Location. Germany, Hamburg, river Elbe (“Elbe bei Hamburg”).
Remarks from the catalogue. Collected by A. Westphalen (no date given). Received by the museum on April 26, 1900. Determined by Heinze in 1935 as Gordius mülleri.
Occurs in publications.
Remarks. The cuticle does not contain areoles (Fig.
Material examined. V5523 (1 female). Fig.
Location. Germany, Rahlstedt (“Alt-Rahlstedt”), under a flower pot.
Remarks from the catalogue. Collected by Ilse Schultze (date not given). Received by the museum on September 15, 1900. Determined by Heinze in 1935/36 as “Gordius ? aquaticus”.
Occurs in publications.
Remarks. The cuticle of this specimen is smooth (Fig.
Material examined. V5773 (1 male). Fig.
Location. Germany, Eppendorfer Mühlenteich, Hamburg.
Remarks from the catalogue. Collected by “Lehrer” (teacher ?) Wagner (date unknown). Received by the museum on July 2, 1901. Determined by Heinze in 1935 as Gordius mülleri (= muelleri).
Occurs in publications.
Remarks. The body cuticle is heavily covered with some artificial crust (Fig.
Material examined. V6341 (1 female). Fig.
Location. Germany, Hessen, Stream Olbe.
Remarks from the catalogue. Collected by G. Ulmer in July 1900. Received November 14, 1903. Determined by Heinze in 1935 as Gordius stellatus.
Occurs in publications.
Remarks. The cuticle is smooth with some irregularly placed shallow elevations (Fig.
Material examined. V6488 (1 male). Fig.
Location. Germany, Hannover, River Örtze.
Remarks from the catalogue. Collected by Th. Lindemann (no date given). Received by the museum on July 28, 1904. Determined by Michaelsen in 1904 as Gordius aquaticus and by Heinze in 1935 as Gordius setiger.
Occurs in publications.
Remarks. The cuticle has very indistinct elevations that might represent areoles (Fig.
Material examined. V6826 (2 males and one specimen of unknown sex, posterior end is missing). Fig.
Location. China, province Fokien, Futschou.
Remarks from the catalogue. Collected by “Cons. G. Siemssen” (no date given). Received by the museum on April 4, 1906.
Occurs in publications. unknown
Remarks. The specimen has a smooth cuticle (Fig.
Material examined. V7456 (1 female). Fig.
Location. China, province Fujian (as “Prov. Fo-Kien”).
Remarks from the catalogue. collected by “Consul G. Siemssen” (no date given), received by the museum on September 8, 1911. Determined as “Gordius” by Angener or Augener (as “Ang. det. 1931” or “Aug. det. 1931”).
Occurs in publications. unknown
Remarks. The specimen has a smooth body cuticle (Fig.
Material examined. V7724 (1 male). Fig.
Location. China, province Fujian (as “Prov. Fokien”).
Remarks from the catalogue. collected by “Consul G. Siemssen” (no date given), received by the museum on September 4, 1912. Determined as “Gordiide”.
Occurs in publications. unknown
Remarks. The body cuticle is smooth (Fig.
Material examined. V8077 (1 male). Fig.
Location. Italy, Sorgono, Sardinia.
Remarks from the catalogue. collected by A.H. Krause (no date given), received by the museum on March 14, 1913. Determined as “Gordius”.
Occurs in publications. unknown
Remarks. The cuticle is not in good shape, but it appears to be smooth (Fig.
Material examined. V9075 (2 females). Fig.
Location. France, surroundings of Montpellier.
Remarks from the catalogue. collected by K. Schmalfuss (no date given). Received by the museum on December 23, 1920. Originally determined as “Gordiide”. Although the catalogue indicates only one specimen, two females were found in the vial, named here (I) and (II).
Occurs in publications. unknown
Remarks. Both specimens have a smooth cuticle covered by some dirt (Fig.
Material examined. V9214 (unknown sex, because posterior end is missing). Fig.
Location. Chile, Victoria in province Malleco (swimming in river).
Remarks from the catalogue. collected by C. Bock in February 1920. Received by the museum on August 12, 1921.
Occurs in publications. unknown
Remarks. The cuticle is smooth (Fig.
Material examined. V10154 (1 male). Fig.
Location. Germany, Hamburg, Süderelbe (river Elbe).
Remarks from the catalogue. Collected by N. Peters (date only given), received by the museum on June 7, 1927. Determined by Heinze in 1935 as Gordius muelleri (as mülleri).
Occurs in publications. unknown.
Remarks. Unfortunately the posterior end of this specimen was lost during preparation for SEM, but it was a male posterior end with a postcloacal crescent. The cuticle is smooth (Fig.
Material examined. V10529 (1 male). Fig.
Location. Brazil, Itatiaia, state of Rio de Janeiro, 700m, in a puddle.
Remarks from the catalogue. Collected by F. Ohaus (date unknown), donated to the museum by Titschak, received by the museum on March 15, 1928.
Occurs in publications. unknown
Remarks. This specimen clearly belongs to the genus Gordius due to the presence of a postcloacal crescent (Fig.
Material examined. V10599 (1 male). Fig.
Location. probably Croatia, Rijeka. Writing in catalogue: “Umgeg. v. Fiume”, Fiume is the Italian and Hungarian name for Rijeka (source: Wikipedia). This is supported by the added word “Jugoslavien” (Yugoslavia) in different handwriting.
Remarks from the catalogue. no collector indicated, given to the museum by de Gries. Received by the museum on September 24, 1928. Additional remark: from Lacerta muralis (common wall lizard).
Occurs in publications. unknown
Remarks. This is a Gordius specimen with smooth cuticle (Fig.
Material examined. V10955 (1 male). Fig.
Location. Germany, stream Linau in Witzeeze, Schleswig-Holstein.
Remarks from the catalogue. Collected by G. Duncker (date only given on label in vial: August 26, 1929), received by the museum on September 2, 1929. Determined by Heinze in 1935 as Gordius setiger. (label in vial says: “Gordius spec. setiger oder mülleri. Vermutl. doch setiger Schneider”).
Occurs in publications.
Remarks. Unfortunately the posterior end of this specimen was lost during preparation for SEM, but it was a male posterior end with a postcloacal crescent. There are flat areoles (Fig.
Material examined. V11466 (1 male and fragments of probably 2 further specimens; only male was investigated). Fig.
Location. Brazil, Boiteuxburgo, Santa Catarina.
Remarks from the catalogue. Collected by Paul Missfeldt (no date given). Received by the museum on December 5, 1930. Originally determined as “Gordius”.
Occurs in publications. unknown
Remarks. The body cuticle is smooth (Fig.
Material examined. V11943 (1 male). Fig.
Location. Hungary. No further specification is given here, but number V11942 is also listed in the catalogue as a gordiid, with the same collector and location “Ungarn, Warmbad Villach, Thermenabfluss”. This means the worm was found in the drainage of a spa. Today, Villach is a village in Austria, until 1918 Austria and Hungary were united, which could explain the catalogue entry. It is assumed that the same locality is valid also for V11943. A vial V11942 is present in the collection, but it contains no specimen.
Remarks from the catalogue. Collected by Gebhard (no date given) and received by the museum on September 28, 1933. Original determination as “Gordiiden”.
Occurs in publications. unknown
Remarks. The specimen has a postcloacal crescent bordering the point of bifurcation of the tail lobes (Fig.
Material examined. V12712 (1 female). Fig.
Location. Germany, Schiffbeker Moor, Hamburg.
Remarks from the catalogue. Collected by Mrs. Helme (date unknown) and received by the museum on October 12, 1937.
Occurs in publications. unknown
Remarks. The cuticle is smooth (Fig.
Material examined. V13285 (1 male)
Location. Japan, Katsura river, Arashiyama, Kyoto prefecture (35°00'48"N, 135°40'40"E).
Remarks from the catalogue. Collected by Kazuo Isobe on March 13, 2008.
Occurs in publications.
Remarks. This male Gordius specimen could not be determined further, see
Material examined. V13305 (1 male)
Location. Japan, Kumano, Misen.
Remarks from the catalogue. Collected by Takuya Sato on September 14, 2009. Received by the museum in 2011.
Occurs in publications.
Remarks. Specimen is a voucher for a phylogenetic analysis in
Material examined. V13306 (1 male)
Location. Japan, Kumano, Nadani.
Remarks from the catalogue. Collected by Takuya Sato on October 14, 2007. Received by the museum in 2011.
Occurs in publications.
Remarks. Specimen is a voucher for a phylogenetic analysis in
Material examined. V13307 (1 male)
Location. Japan, Arida, Ninomata.
Remarks from the catalogue. Collected by Takuya Sato on September 30, 2009. Received by the museum in 2011.
Occurs in publications.
Remarks. Specimen is a voucher for a phylogenetic analysis in
Material examined. V13308 (1 male)
Location. Japan, Arida, Ninomata.
Remarks from the catalogue. Collected by Takuya Sato on September 13, 2009. Received by the museum in 2011.
Occurs in publications.
Remarks. Specimen is a voucher for a phylogenetic analysis in
Material examined. V13309 (1 male)
Location. Japan, Kumano, Nadani.
Remarks from the catalogue. Collected by Takuya Sato on November 10, 2008. Received by the museum in 2011.
Occurs in publications.
Remarks. Specimen is a voucher for a phylogenetic analysis in
Material examined. V13310 (1 male)
Location. Japan, Kumano, Nadani.
Remarks from the catalogue. Collected by Takuya Sato on October 15, 2007. Received by the museum in 2011.
Occurs in publications.
Remarks. Specimen is a voucher for a phylogenetic analysis in
Material examined. V13311 (1 male)
Location. Japan, Kumano, Nadani.
Remarks from the catalogue. Collected by Takuya Sato on November 15, 2006. Received by the museum in 2011.
Occurs in publications.
Remarks. Specimen is a voucher for a phylogenetic analysis in
Gordius aquaticus Linné, 1758
Material examined. V12685 (1 male). Fig.
Location. Germany, Hamburg.
Remarks from the catalogue. Collected by M. Grosse (date not given). Received by the museum on August 5, 1937.
Occurs in publications. unknown.
Remarks. The cuticle of this specimen is smooth (Fig.
Gordius balcanicus Schmidt-Rhaesa, 2010
Material examined. V13270 (1 male)
Location. Serbia, Kučaj Mountains, Dubašnica, Mikuljska River.
Remarks from the catalogue. Collected by Ajtić on October 3, 2006.
Occurs in publications.
Remarks. Holotype of this species. See
Gordius balticus Schmidt-Rhaesa, 2010
Material examined. V13268 (1 male)
Location. Estonia, South Estonia (exact location unknown), in rainwater barrel.
Remarks from the catalogue. The collector according to the label is “Elli” in 2008.
Occurs in publications.
Remarks. Holotype of this species. See
Material examined. V13288 (1 female)
Location. Estonia, rainwater container in Suurküla, Häädemeeste.
Remarks from the catalogue. Collected by M. Looring on August 23, 2006.
Occurs in publications.
Remarks. Paratype of this species. See
Gordius digitatus Schmidt-Rhaesa, 2010
Material examined. V13271 (1 male)
Location. Croatia, Plitvice Lakes (Plitvička Jezera).
Remarks from the catalogue. Collected by M. Teiwes on August 8, 2000.
Occurs in publications.
Remarks. Holotype of this species. See
Gordius helveticus Schmidt-Rhaesa, 2010
Material examined. V13272 (1 male)
Location. Swizerland, Meiringen (Berner Oberland), altitude 793m.
Remarks from the catalogue. Collected by Axel Groenveld on July 18, 2008.
Occurs in publications.
Remarks. Holotype of this species. See
Material examined. V13277 (1 male)
Same data as for V13272, paratype of this species. See
Gordius karwendeli Schmidt-Rhaesa, 2010
Material examined. V13273 (1 male)
Location. Germany, Karwendel Mountains, 4–4.5 km NE of Mittenwald. Found in a (not water covered) gravel-bed along stream Seinsbach at roadside during heavy rains.
Remarks from the catalogue. Collected by Maaike van Rijn and Axel Groenveld on June 25, 2002.
Occurs in publications.
Remarks. Holotype of this species. See
Gordius longissimus Römer, 1895
Material examined. V2229 (1 female). Fig.
Location. "Südsee" (= South Sea), no further specification.
Remarks from the catalogue. Collected by “M.G.” (writing not explicit). No collection date given and no date when received by the museum. From Saturnia (Lepidoptera). Determined by Römer in 1893 as G. longissimus, confirmed by Heinze in 1935.
Occurs in publications.
Remarks. Holotype of this species.
Gordius paranensis Camerano, 1892
Material examined. V5035 (1 male, 2 females). Fig.
Location. Paraguay, San Bernardino.
Remarks from the catalogue. Collected by “Cons. Wiengreen” (no date given). Received by the museum on April 30, 1898. Determined as “Gordius”.
Occurs in publications. unknown
Remarks. The male contains in the posterior end a semicircular row of bristles anterior of the cloacal opening (Fig.
Material examined. V11709 (8 males; 1 male investigated by SEM). Fig.
Location. Brazil, Boiteuxburgo, Santa Catarina.
Remarks from the catalogue. Collected by Paul Missfeldt (date unknown). Received by the museum on September 8, 1932. Originally determined as “Gordiide”.
Occurs in publications. unknown
Remarks. The male contains in the posterior end the semicircular row of bristles characteristic for this species (Fig.
Gordius pesici Schmidt-Rhaesa, 2010
Material examined. V13274 (1 male)
Location. Montenegro, Budva.
Remarks from the catalogue. Collected by Vladimir Pešić on September 2, 2006.
Occurs in publications.
Remarks. Holotype of this species. See
Gordius plicatissimus Heinze, 1952
Material examined. V5524 (1 male). Fig.
Location. Swizerland, Grindelwald.
Remarks from the catalogue. Collected by Michaelsen (no date given). Received by the museum on September 15, 1900. Determined as “Gordius”. Host: Decticus verrucivorum.
Occurs in publications. unknown
Remarks. Areoles are large, prominent and slightly oval (Fig.
Material examined. V13332 (1 male). Fig.
Location. Austria, Salzburg, St. Gilgen, Falkenstein, 717 m. The specimen was found alive during excavations of a historical source in about 2 m depth.
Remarks from the catalogue. Collected by an unknown person on August 30, 2011. Donated to the museum by Robert Lindner, Haus der Natur in Salzburg, Austria in July 2012.
Occurs in publications. no
Remarks. Areoles are prominent and slightly oval, the arrangement is perpendicular to the longitudinal body axis (Fig.
Gordius cf. polychaetus Tang, 1934
Material examined. V13283 and V13284 (1 male each)
Location. Japan, Sobu Valley, Kaminyu, Yogo Town, Shiga Prefecture (35°34'14"N, 136°12'41"E), on the surface of snow.
Remarks from the catalogue. Collected by Daiki Kado on November 22, 2008.
Occurs in publications.
Remarks. Both specimens have numerous fine bristles on their cuticle, which corresponds to the description of G. polychaetus Tang, 1934. See
Gordius serratus Schmidt-Rhaesa, 2010
Material examined. V13280 (1 male)
Location. Papua New Guinea, Lahamenegu piggery, Goroka, Eastern Highlands Province, from river water.
Remarks from the catalogue. Collected by Ifor Owen, presumably in 1975.
Occurs in publications.
Remarks. Holotype of this species. See
Material examined. V13281 (1 male)
Same data as for V13280, paratype of this species. See
Gordius setiger Schneider, 1866
Material examined. V2224 (1 male). Fig.
Location. Germany, Hamburg, Alster.
Remarks from the catalogue. Collected by Graeser (no collection date or date when received by the museum given). Determined by Römer in 1893 as Gordius violaceus and by Heinze in 1935 as Gordius setiger.
Occurs in publications.
Remarks. The SEM reinvestigation shows inconspicuous areoles (Fig.
Material examined. V5940 (1 male). Fig.
Location. Germany, river Elbe close to Lauenburg.
Remarks from the catalogue. Collected by “Wilh. Meyer” on June 1, 1902. Received by the museum on July 1, 1902. Determined by Heinze in 1935 as Gordius mülleri.
Occurs in publications.
Remarks. Areoles on the cuticle are roughly polygonal in shape and attach closely to each other (Fig.
Gordius cf. setiger Schneider, 1866
Material examined. V9000 (1 male). Fig.
Location. Germany, lake “Behler See” near Timmendorf, Holstein.
Remarks from the catalogue. collected by “Gast” with unknown date. Received by the museum on June 15, 1920. Determined by Heinze in 1935 as Gordius setiger.
Occurs in publications.
Remarks. The two species G. setiger and G. albopunctatus are very similar and differ only in few details. According to
Material examined. V9900 (2 males). Fig.
Location. Germany, Hamburg, river Elbe, Parkhafen (writing is not explicit).
Remarks from the catalogue. collected by H. Meves (date not given). Received by the museum on May 29, 1925. Originally determined as “Gordiiden”. On the vial label, but not in the catalogue is written: determined as Gordius setiger by R. Arndt in 1968.
Occurs in publications. unknown
Remarks. The two males of this lot show characters as described above for G. setiger and G. albopunctatus (see above under V9000). Male I has polygonal areoles (Fig.
Gordius spiridonovi Schmidt-Rhaesa, 2010
Material examined. V13286 (1 male)
Location. Estonia, Lake Peipsi (about 58°40'N, 27°30'E).
Remarks from the catalogue. Collected by an unknown person on June 9, 1980.
Occurs in publications.
Remarks. Holotype of this species. See
Material examined. V13287 (1 male)
Location. Russia, Udoha River (tributary of the Shelon River), near Borovit, Pskov Region.
Remarks from the catalogue. Collected by T. Timm on July 7, 1971.
Occurs in publications.
Remarks. Paratype of this species. See
Gordius terminosetosus Schmidt-Rhaesa, 2010
Material examined. V13279 (1 male)
Location. Estonia, Sauga Stream, western Estonia (58°31.54'N, 24°42.16'E).
Remarks from the catalogue. Collected by H. Timm on August 16, 1993.
Occurs in publications.
Remarks. Holotype of this species. See
Gordius undulatus Heinze, 1937
Material examined. V5521 (1 male). Fig.
Location. Germany, Hamburg, Winterhude.
Remarks from the catalogue. Collected by Braun (writing not explicit, but this name is given by
Occurs in publications.
Remarks. This is the holotype for this species. (
Gordius zwicki Schmidt-Rhaesa, 2010
Material examined. V13275 (1 male)
Location. Russia, Russian Far East, small forest tributary in the upper course of Milogradovka River, 25 km NW of Milogradovo.
Remarks from the catalogue. Collected by Peter Zwick on June 13, 1998.
Occurs in publications.
Remarks. Holotype of this species. See
Material examined. V13276 (1 male)
Same data as for V 13275, paratype of this species. See
Nectonema is the only marine genus of horsehair worms (Nematomorpha) and includes 5 species. Two of them (N. agile, N. munidae) were repeatedly found, all other species were only caught on single occasions.
Nectonema munidae Brinkmann, 1930
Material examined. V13289 (2 females)
Location. Norway, Korsfjorden near Bergen, 550-700m.
Remarks from the catalogue. Collected by A. Schmidt-Rhaesa, Martin V. Sørensen, Reinhardt Møbjerg Kristensen and Henrik Glenner in August 2009. From host Munida tenuimana.
Occurs in publications. none.
Remarks. none.
Material examined. V13290 (2 males)
Location. Norway, Korsfjorden near Bergen, 550-700m.
Remarks from the catalogue. Collected by A. Schmidt-Rhaesa, Martin V. Sørensen, Reinhardt Møbjerg Kristensen and Henrik Glenner in August 2009. From host Munida tenuimana.
Occurs in publications. none.
Remarks. none.
Species in this genus have two types of areoles, the second, larger type are large areoles that seem to be fusion products of several smaller areoles (
Parachordodes tolosanus (Dujardin, 1842)
Material examined. V2220 (3 males and 2 females are indicated in the catalog, but actually only 2 males and 1 female are present; female transferred to new catalogue number, see below). Fig.
Location. Göttingen, Hainholz, in source (Germany).
Remarks from the catalogue. Collected by Schlotthauber (no collection date or date when received by the museum given). Determined by Römer in 1893 as Gordius violaceus, corrected by Heinze in 1935 to Parachordodes tolosanus.
Occurs in publications.
Remarks. Collector and collection dates are similar to V2218 determined as Gordionus wolterstorffii. The fate of two missing specimens is unknown. Two specimens (1 male and 1 female) were reinvestigated by SEM. The cuticle of the male contains superareoles (Fig.
Material examined. V2236 (I) (1 male) One specimen is indicated in the catalogue, but 2 specimens are present. The second specimen is determined as Gordionus sp.; Fig.
Location. Göttingen, Germany
Remarks from the catalogue. Collected by Schlotthauber (no collection date or date when received by the museum given). Determined by Schlotthauber in 1854 as Gordius aquaticus, corrected by Heinze in 1935 to Parachordodes tolosanus. Host: Harpalus ruficornis crossed out and replaced by Ophonus cephalotes.
Occurs in publications.
Remarks. Superareoles are present (Fig.
Material examined. V2237 (1 male). Fig.
Location. Göttingen, Germany
Remarks from the catalogue. Collected by Schlotthauber (no collection date or date when received by the museum given). Determined by Schlotthauber in 1854 as Gordius aquaticus, corrected by Heinze in 1935 to Parachordodes speciosus. Host: Harpalus distinguendus.
Occurs in publications.
Remarks.SEM reinvestigation shows superareoles on the cuticle (Fig.
The genus Paragordionus resembles Gordionus in many respects, but the cuticle contains two types of areoles. The larger areoles are clustered together, such clusters can be composed of two or of several areoles. The large areoles are called megareoles (see
Paragordionus dispar (Müller, 1926)
Material examined. V7449 (II) (1 male). Fig.
Location. Germany, Berchtesgaden.
Remarks from the catalogue. Collected by “Heinr. Baur” (not well readable) (no date given). Received by the museum on August 1, 1911. Determined by Heinze in 1935 as Paragordionus dispar. Although the catalogue says that only one specimen belongs to this lot, two specimens are present. These are marked with (I) and (II).
Occurs in publications.
Remarks. The two specimens (I) and (II) both belong to the genus Paragordionus, because they possess clusters of megareoles (see
Paragordionus ibericus Schmidt-Rhaesa & Cieslak, 2008
Material examined. V13260 (1 male)
Location. Spain, Puerto de los Cotos, Arroyo de la Laguna Grande de Penalara (40°49'59"N, 3°56'09"W), 1600 m.
Remarks from the catalogue. Collected by I. Ribera and A. Cieslak on July 1, 2006. Received by the museum on July 3, 2008.
Occurs in publications.
Remarks. Holotype of this species. See
Material examined. V13261 (1 male)
Same data as V13260 (but no type material), see there.
Paragordionus vejdovskyi (Janda, 1894)
Material examined. V7449 (I) (1 male). Fig.
Location. Germany, Berchtesgaden.
Remarks. See V7449 (II) under Paragordionus dispar for all remarks and explanations.
Specimens belonging to this genus are easy to recognize, because females have three lobes around the central and terminal cloacal opening and males have very long paired tail lobes (length: width 2:1 or longer). This genus includes the only species among Nematomorpha, where parthenogenetic reproduction was shown (P. obamai;
Paragordius sp.
Material examined. V3039 (2 males). Fig.
Location. Mexico, Hacienda de Ixtapa Tepic.
Remarks from the catalogue. Collected by Dr. P. (?) Hacker (no date given). Received by the museum on October 2, 1908. Determined as “Gordiide”. Only 1 male is registered in the catalogue, while 2 males were found in the vial.
Occurs in publications. unknown
Remarks. The long tail lobes of the male posterior end are characteristic for the genus Paragordius (Fig.
Material examined. V8948 (1 female). Fig.
Location. Chile, river Estero Manco (as “Fluss Manco”) near Coronel.
Remarks from the catalogue. Collected by R. Paessler on December 16, 1915, received by the museum on January 15, 1920. Originally determined as “Gordiide”.
Occurs in publications. unknown
Remarks. The posterior end has three lobes, two broader and a slender one, which is typical for females of the genus Paragordius (Fig.
Material examined. V8962 (1 female). Fig.
Location. Chile, river Estero Manco (as “Fluss Manco”) near Coronel.
Remarks from the catalogue. Collected by R. Paessler in January 1917, received by the museum on January 15, 1920. Originally determined as “Gordiide”.
Occurs in publications. unknown
Remarks. The posterior end has three lobes, two broader and a slender one, which is typical for females of the genus Paragordius (Fig.
Paragordius obamai Hanelt, Bolek & Schmidt-Rhaesa, 2012
Material examined. V13265-13267 (1 female each number)
Location. Reared from a cyst from a Biomphalaria pfeifferi snail collected at Kasabong stream, a small stream in Nyanza province, Kenya (approx. 1170 m altitude), which flows into Winam Bay of Lake Victoria, 2 km to the south.
Remarks from the catalogue. Received from Ben Hanelt.
Occurs in publications.
Remarks. V13265 is the holotype for this species, V13266 and V13267 are paratypes. See
Pseudochordodes is a genus with distribution from the southwestern Nearctic to the Neotropis. Eight species have been described. The males have an unbranched posterior end (= without tail lobes) and the cuticle of both sexes has two types of areoles (see
Pseudochordodes meridionalis Carvalho & Feio, 1950
Material examined. V2930 (1 female). Fig.
Location. Guatemala
Remarks from the catalogue. From bequest of “O. Hoffmann der Jul. Groth”. Received by the museum on February 14, 1908. Originally determined as “Chordodes”.
Occurs in publications. unknown
Remarks.SEM reinvestigation reveals larger and smaller areoles (Fig.
Material examined. V2226 (1 female). Fig.
Location. USA, Arizona, Prescott.
Remarks from the catalogue. Collected by Rodenburg (written in
Occurs in publications.
Remarks. The cuticle of this specimen shows areoles with no clear border and no interareolar structures (Fig.
Material examined. V2233 (1 male). Fig.
Location. “Bahia”
Remarks from the catalogue. Collected by Bolau (no date given). Received by the museum on September 23, 1893. Determined by Römer in 1893 as C. variopapillatus, holotype of this species. Heinze in 1935 writes “specimen in bad condition, not to determine [sehr schlecht erhalten, indeterminable]” and replaces “C. variopapillatus” by “Gordius spec.?”.
Occurs in publications.
Remarks. The specimen is indeed in bad condition. No areoles were observed on the cuticle (Fig.
Material examined. V5522 (1 specimen, sex unknown). Fig.
Location. Germany, Ahrensburg.
Remarks from the catalogue. Collected by Post (no date given). Received by the museum on September 8, 1900. Determined by Heinze in 1935 as “Gordius ? undulatus”.
Occurs in publications. unknown (not in
Remarks. The posterior end is damaged, the sex cannot be determined. The cuticle is heavily covered with some covering, probably fungus (Fig.
Material examined. V6622 (sex not determined). Fig.
Location. Germany, Hamburg.
Remarks from the catalogue. Collected by Rump (no date given). Received by the museum on November 10, 1904. Host: Vanessa io (European peacock, currently Aglais io or Inarchis io, Lepidoptera).
Occurs in publications. unknown
Remarks. The cuticle is smooth and has some folds (Fig.
Material examined. V7028 (1 specimen). Fig.
Location. Portugal, Leixões
Remarks from the catalogue. Collected by F. Ohaus on August 25, 1904. Received by the museum on January 1, 1907. Determined as “Gordiide”.
Occurs in publications. unknown
Remarks. Cuticle smooth, covered with dirt or fungi (Fig.
Specimens of Gordius sp., ventral view of posterior ends and cuticle in midbody sections. The specimen number is indicated in the lower left of each image. A, B, G. Posterior ends with postcloacal crescent (pcc) and cloacal opening (co). The cuticular surface (C–F, H) is in some cases covered with encrusting material (C) or dirt (F, H). Diagonal lines are created by muscular fibers underlying the cuticle.
Specimens of Gordius sp., ventral view of posterior ends and cuticle in midbody sections. The specimen number is indicated in the lower left of each image. A–E, G–I. Cuticular surface. This is either smooth (A–C, G), has keel-like structures (D) or areoles (E, H, I). F, J, K. Posterior ends with postcloacal crescent (pcc) and cloacal opening (co). The posterior end of specimen V4611 has a fringed rim around the cloacal opening (J, K).
Specimens of Gordius sp., ventral view of posterior ends and cuticle in midbody sections. The specimen number is indicated in the lower left of each image. A, D, E, G, H, J–L. Cuticular surface, see text for explanation. B, F, I, M. Posterior ends with postcloacal crescent and cloacal opening (co). C. Magnification of short bristles on the tail lobes of V4981.
Specimens of Gordius sp., ventral view of posterior ends and cuticle in midbody sections. The specimen number is indicated in the lower left of each image. A, C, D, G, H–K, M. Cuticular surface, see text for explanation. B, E, F, L, N. Posterior ends with postcloacal crescent and cloacal opening (co). O. Areoles on the tail lobes of V10599.
A–F. Specimens of Gordius sp., ventral view of posterior ends and cuticle in midbody sections. G, H. Gordius aquaticus (V12685), cuticle (G) and posterior end (H). I. Cuticular surface of G. longissimus (V2229). J, K. Cuticle (J) and posterior end (K) of G. paranensis (V5035). Note row of bristles on the posterior end (arrows in K). L. Posterior end of G. paranensis (V11709), arrows as in K.M–O. Cuticle (M, N) and posterior end (O) of G. plicatissimus. The specimen number is indicated in the lower left of each image.
A, B. Cuticle (A) and posterior end (B) of Gordius undulatus (V5521). C–J.Acutogordius cf. acuminatus, specimens V2213 (C–E), V7351b (F, G) and V11465 (H–J). Note longitudinal crests or keels on tail lobes of V11465 (arrows in J) and scattered bristle-like structures on the cuticle (circles in I) of the same specimen (see text).
A–H. Paragordius sp., details of cuticle (A, B, E, G) and shape of the posterior ends of males (C, D; V3039) and females (F; V8948 and H; V8962). Arrow in F and H indicates the dorsal lobe. I–K. Beatogordius sp., longitudinal row of areoles (I), magnification of areoles (J) and posterior end (K).
A–D. Beatogordius sp., cuticular pattern (A), detail of the cloacal opening (B), overview (C) and detail (D) of the posterior end, showing the cloacal opening (co) and the postcloacal spines (pcs). E–O. Gordionus sp., cuticular pattern and posterior ends of specimens V2215, V2217, V2236 (II), two specimens of V9615 (I and II) and V11117. H shows detail of precloacal bristles anterior to the cloacal opening (co). N shows precloacal bristles, O shows the cloacal opening (co) and some postcloacal spines (pcs).
A–C. Cuticular surface of three specimens of Gordionus sp. (V12470, V13361, V13362). D–H. Gordionus scaber. Cuticular surface (D) and posterior end (E) of V2221, posterior end (G), magnification (F) and detail from the precloacal bristles (H) from V10924. I–L. Gordionus violaceus. Cuticular surface (I) and posterior end (J) of V2225, cuticular structure of two females of V11455 (K, L).
A–E. Gordionus wolterstorffii, cuticular surface (A–C) and posterior end (D, E) in specimens V2218, V2222 and V5255. Posterior end with precloacal bristlefields (pbf) and postcloacal spines (pcs). F–H. Paragordionus vejdovskyi, pattern of areoles on the cuticle. F. Overview with ventral midline (vml) and two types of areoles, simple areoles and larger megareoles in clusters (two of these are encircled). G. Maginification from F.H. Large cluster (encircled) of megareoles. I, J. Paragordionus dispar with smaller clusters of megareoles. K–Q. Parachordodes tolosanus. K. Superareoles, which are characteristic for Parachordodes, are encircled. L. Posterior end of V2220 with precloacal bristlefields (pbf), M. Magnification of precloacal bristlefields. N, O. Cuticular structure of V2236I with superareoles. P. Cuticle of V2237, Q. Posterior end of V2237 with precloacal bristlefields (pbf) and cloacal opening (co).
A, B. Pseudochordodes meridionalis, V2930. Cuticular structure in low (A) and higher (B) magnification, encircled in B is a tubercle areole. C–O. Chordodes sp., specimen V2230 (C–G) was formerly designated as holotype of C. pilosus. C. Overview onto the ventral side. Note elevated clusters on both sides of the ventral midline (vml). D. Overview onto the lateral side. Note scattered elevated clusters (arrow indicates one of these clusters). E. Higher magnification shows that the clusters are composed of elevated areoles. F. Clusters of crowned areoles (crar) and circumcluster areoles (ccar) are strongly eroded. Image shows a cluster from next to the ventral midline which includes crowned areoles with long apical filaments. G. Strongly eroded cluster from the lateral body side. H–O. Specimen V5320; H–K female specimen, L–O male specimen. H, I. Arrangement of crowned areole clusters with long filaments next to the ventral midline (vml). J. Two central crowned areoles within a cluster. K. Crowned areoles (crar) surrounded by circumcluster areoles (ccar). L–O. Cuticle and areoles in the male, see female for abbreviations. P–S. Chordodes brasiliensis, different specimens of number V7351. Cuticle shows scattered elevated crowned areoles among simple areoles (sar). Specimen V7351d has part of the cuticle covered by rests of the larval cuticle (lc).
A–F. Chordodes brasiliensis, different specimens of number V7351. A, B. Higher magnification of crowned areoles. C, D. Crowned areoles with long apical filaments along the ventral midline (vml) in female specimen V7351e. E, F. Posterior end of a male (V7351c) with cloacal opening (co) and bristles around and in the region of the opening. G–J. C. jandae or C. skorikovi. G. Cuticular structures on the ventral side showing crowned areoles clusters with long filaments along the ventral midline and adjacent transition to clusters with short filaments (arrow). H, J. Magnification of crowned areoles. I. Different types of areoles: besides simple and crowned areoles (shown but not marked) there are bulging areoles (buar), thorn areoles (thar) and tubercle areoles (tuar). K, L. Chordodes japonensis, cuticular structure with tubercle areoles (tuar, encircled). Part of the cuticle is covered with larval cuticle (lc). M–P.Chordodes cf. japonensis, cuticle with different types of areoles: tubercle areoles (tuar, encircled in M), thorn areoles (thar) and circumcluster areoles (ccar) surrounding crowned areoles. Posterior end with cloacal opening (co).
A–F.Chordodes cf. joyeuxi. A. Female posterior end with terminal cloacal opening. B. Overview on cuticle with different types of areoles: tubercle areoles (tuar), thorn areoles (thar) and cluster of crowned and circumcluster areoles (crcl) (here a cluster with long filaments is present). C. Magnification of thorn areoles. E Magnification of tubercle areoles. D, F. Clusters of central crowned areoles and circumcluster areoles (ccar). G–I. Chordodes polycoronatus. G. Abundant large crowned areole clusters (one cluster is encircled). H. Posterior end with precloacal bristlefields (pbf). I. Magnification showing similar structure of areoles in the clusters.
Chordodes jelkae sp. n.; A–E from holotype (V10960I). A. Overview on the cuticle from the lateral side showing elevated crowned areole clusters. B. Overview of the cuticle from the ventral side showing clusters with crowned areoles with long filaments along the ventral midline (vml). C. Fine structure of simple areoles (sar), crowned areoles (crar) and circumcluster areoles (ccar). Some areoles may be either type (*). D. Magnification of simple areoles. E. Arrangement of simple areoles (sar) and structure of tubercle areoles (tuar). F. Cuticle from second specimen (V10960II).
Two persons have helped very much during this project. Vladimira Oldenburg made most of the SEM investigation during a student project. Helma Roggenbuck took care of handling the collection and updating everything according to new results. Many thanks to both of them.