163brachiopoda groenlandica

Steenstrupia Thomsen, E. Brachiopoda Groenlandica – an historical and taxonomic review of brachiopods from Greenland. – Steenstrupia 32 (2): 163–186. Copenhagen, Denmark. December 2012. ISSN 0375-2909.

A review of the collection history of brachiopods from Greenlandic waters demonstrates the impor-tance of international and national expeditions during the period 1860–1992. Systematic studies of the brachiopod collection at the Natural History Museum of Denmark, University of Copenhagen, reveal seven species from about 150 localities. The species are: Pelagodiscus atlanticus (King, 1868) – very sparse, Cryptopora gnomon Jeffreys, 1869 – very sparse, Hemithiris psittacea (Gmelin, 1791) – very abundant, Arctosia arctica (Friele, 1878) – very sparse, Terebratulina septentrionalis (Couthouy, 1838) – abundant, Glaciarcula spitzbergensis (Davidson, 1852) – sparse, and Macandrevia cranium (Müller, 1776) – sparse. The collection was found to house the syntypes of Hemithiris psittacea. The geographical distribution of the different species is related to bathymetric depth, temperature, currents and water masses. There is a general preference for depths less than 300 m, with the exceptions of P. atlanticus and C. gnomon, both of which occur at abyssal depths, and T. septentrionalis, which has a wider range (100–600 m). The latter is common along the southern part of the west coast and in the so-called "Atlantic" fjords that are influenced by warm Irmiger Water. M. cranium prefers warm At-lantic Water (Thomsen 1990) and is therefore restricted to the southern part of the west coast and other areas of Irmiger Water from the Irmiger Current. A. arctica and G. spitzbergensis prefer colder water, which is reflected in their occurrence, the former in areas with Polar Water from the East Greenland Current and the latter in areas influenced by the temperate water of the West Greenland Current. The most abundant and frequent species is H. psittacea, a circumpolar species that prefers cold water. It is found all along the coast of Greenland, including the fjords.

Keywords: Brachiopoda, Greenland, expeditions, taxonomy, distribution

Elsebeth Thomsen: Tromsø University Museum, Department of Natural Sciences, NO-9037 Tromsø, Norway. E-mail: elsebeth.thomsen@uit.no

Steenstrupia 32 (2): 163–186.

Brachiopoda Groenlandica – an historical and taxonomic review of brachiopods from GreenlandELSEBETH THOMSEN

INTRODUCTION

Brachiopods from the Greenland area have been collected since the first record was pub-lished in 1785. The Zoological Museum of the University of Copenhagen (ZMUC), now part of the Natural History Museum of Denmark (SNM), possesses a collection comprising dried and ethanol preserved material from about 150 localities (Appendix 1 gives named localities). The specimens have mainly been recovered by Danish Expeditions during the late 19th Century and the 20th Century.

The aim of the present study is to present 1) an historical review of the recovery of brachiopods from Greenlandic waters and 2) a taxonomic part based on the collection of ZMUC, listing

Greenlandic brachiopod references. Brachiopods from Greenland in collections situated in other institutions have not been studied.

PREVIOUS INVESTIGATIONS

"von den Kalten Ufern…"The first record of brachiopods from Greenlandic waters was published in 1785 by Johann Hiero-nymus Chemnitz in volume eight of his monu-mental work Neues systematisches Conchylien-Cabinet. Here we find several brachiopod species listed as Anomia species, e.g., Anomia rostrum psittaci, the bird or parrot beak anomia (his table

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78, fig. 713a–c; see Fig. 1). Chemnitz stated that he had six specimens of this rare "Gattung", "die ich einst unvermuthet aus der Strasse Davids, von den Kalten Ufern des westlichen Grønlandes bekommen, daher ich sie ab wenigsten erwartet hätte". Chemnitz' figures depict a complete speci-men (a), a pedicle valve (b) and a brachial valve (c), both of the latter showing the interior. The specimens were "Ex Museo nostro", i.e., they belonged to Chemnitz' own collection and were the first illustrated specimens of Hemithiris psit-tacea from Greenland.

Many of the specimens in Chemnitz' book, however, belonged to Lorentz Spengler's private collection and are noted by Chemnitz as being "Ex Museo Spengleriano". Spengler became superintendent for the Royal Kunstkammer in 1771 and sold his private collection to the King in 1814, which is why the specimens are now at SNM/ZMUC.

The present study of the brachiopod material from Greenland revealed a glass with dried speci-

mens of Hemithiris psittacea, with an original label by Spengler, referring to the above men-tioned figures in Chemnitz, see Fig. 1. The loca-tion is given as "Aus Grønland". It is reasonable to assume that Spengler, who was a good friend of Chemnitz, was the provider of his specimens, which is also indicated by the number 13 in the glass. The plate in Chemnitz has the heading "13) Anomien". Thus, the surviving specimens at SNM belonged to Chemnitz collection in 1785. The dried brachiopods later became part of the collection of King Christian VIII and now comprise one pedicle valve, one juvenile pedicle valve, two brachial valves and a tiny pebble, which may have served as a substrate. The speci-mens must have been sampled before 1785 and are the oldest from Greenland.

In 1791 (not 1790, see Kabat & Petit 1988), Gmelin published Linné's Systema Naturae (13th ed.), with annotation by himself. On p. 3348, Anomia psittacea is described with a testa cor-nea (Spengler's label identified the specimens as

Fig. 1. Specimens of Hemithiris psittacea (ZMUC-BRA-1 to -4) from Greenland with the original label by Lorentz Spengler. Specimen no. 13, no. 713a–c in Chr. VIII's collection. From the Chemnitz collection at SNM/ZMUC, i.e., collected before 1785. The book is Chemnitz 1785. Photo by E. Thomsen, Tromsø University Museum (TMU).

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Terebratulina Cornea). Gmelin also presented a synonymy, the oldest reference being Lister (1687), followed by d'Argenville (1742) and Klein (1753) before Spengler and Chemnitz (1785). The habitat is "mari Groenlandiam", the occurrence is very rare. As the Spengler/Chemnitz material was from Greenland, it is not unexpected that SNM's material houses the syntypes. The search for the holotype of this species has been unsuccessful. However, at least two hypotypes from non-Greenlandic localities have been depicted in drawings in Hertlein & Grant IV (1944).

The next record of Greenlandic brachiopods is from Naturhistorisk Tidsskrift (1842−43) by Hans Peter Christian Möller (also spelled Møller) in his work Index Molluscorum Groenlandiæ.Recently, the present author discovered that this paper also exists as a kind of preprint, dated 1842, in the archive at SNM/ZMUC. The preprint is extraordinary, because it also has Möller's per-sonal added notes and fine pencil drawings/sketches. Also found in the archive were addi-tional drawings, some with watercolors.

Schiøtte & Warén (1992) and Schiøtte (1993) published a biography of Möller, who visited Greenland in 1838−40 and collected molluscs assisted by local officials. Möller (1842–43) presented a list of molluscs (including the brachiopod) from West Greenlandic waters. It comprised more than 80 new names, sometimes listed with short, often summary descriptions. Only one species of brachiopod, a specimen of a "Terebratula", "T. psittacea Lmk.", now known as Hemithiris psittacea (Gmelin, 1791), was listed. The preprint was dedicated by hand to Colonibestyrer Kielsen and signed 24 Mars [March] 1843. Möller's personal drawings and notes were no doubt intended to help Kielsen in his continued collecting for Möller, the short printed text not giving sufficient systematic descriptions. Among the drawings is one of the dorsal and lateral exteriors of the brachiopod specimen, "T. psittacea" (Fig. 2). The archive also has a hand-colored drawing and sketches in ink showing the interior of a specimen of the same species (Fig. 3, reference B147E). As ad-ditional help for Kielsen, Möller also marked the listed species in the preprint with marks, such as stars and X's. The latter meant "those species, of

which I do not want any but very excellent, beau-tiful or extraordinarily large specimens". The brachiopod has such an X. Thus, these personal marks also tell us about Möller's interests and pri-orities, e.g., a star meant "those species of which I want all that can be found, large or small, bad or good specimens."

"… of ancestry Mysteriously remote and high" (Jeffreys 1863: 1)Following the collections of Möller, there is a long period before the next Danish publication containing records of Greenlandic brachiopods. Explorations by foreign countries, e.g., Eng-land, Germany and Belgium, became the prime sources of published records. Thus, Jeffreys' British Conchology (1862–65), which dealt with the molluscs, including "this remarkable and peculiar Class Brachiopoda", from the British Isles and surrounding seas, mentioned that "Dr. Wallich has shown me a shell of the last named species [T. cranium] which was brought up by sounding off the east coast of Greenland in 228 fathoms" (Jeffreys 1863: 9). Later (pp. 12−13) this statement is altered to "dead specimens" and to "several depths of 108 and 228 fathoms". Wallich participated in a voyage on board the sloop H.M.S. Bulldog in 1860 in the North Atlantic (The British North Atlantic Telegraph Expedition).

Möbius (1874) listed a sparse brachiopod material collected during the 2. Deutsche Nord-polfahrt 1869−1870 with the H.M.S. Hansa. A specimen of "Terebratula psittacea Gmel." from Jackson Island, East Greenland, and a specimen of "Terebratula cranium Müller" from Shan-non Island (depth 54 m) were included. Both specimens were presented with measurements of length, width and thickness. Measuring the shell was important for Möbius, because he believed that the stability in temperature during the day, which was endured by "nordliche Seetiere", was one of the reasons for their greater size.

In 1875, Jeffreys took part in a cruise with the H.M.S. Valorous to the Davis Strait (and Disko Island) in order to collect zoological samples on the journey back. The purpose was to gain more knowledge of the deep-sea fauna in the North Atlantic and arctic seas (Jeffreys 1876a, b). The Valorous was a supply ship for two other

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ships on the British Arctic Expedition for part of their cruise. The accompanying literature com-prised a manual (Jones 1875) which included a list of Greenland molluscs (including brachio-pods) prepared by "the eminent conchologist of Copenhagen" Otto Andreas Lowson Mörch (Mörch 1875). It lists four species of brachio-pods as Rhynchonella psittacea, Terebratella Spitzbergensis, Terebratulina septentrionalis and Terebratulina cranium. Jeffreys recovered three species on the cruise from Greenlandic waters, including the Davis Strait: T. Spitzbergensis from off Disko Island, R. psittacea from shallow water in the Holsteinsborg area, and a new genus and species, Atretia gnomon (now Cryptopora gno-mon) from deep water in the Davis Strait. Jeffreys took care to note the epifauna on the brachiopod shells as well.

The results of the Valorous were published by Jeffreys (1876a, b), the first paper presenting the new brachiopod genus, Atretia, the last describ-ing three new brachiopod species. In his report on the expedition he mentioned that a "well-equipped Government expedition has within the last few days set out from Bergen with Dr. G.O. Sars in charge". This was the Norwegian North Atlantic Expedition (Den Norske Nordhavs-expedition), which lasted from 1876 to 1878 and included Iceland and Jan Mayen, but no proper Greenlandic waters (Friele 1882, Friele & Grieg 1901). The brachiopods recorded at Jan Mayen were Terebratulina septentrionalis and Terebrat-ula (Liothyris) arctica (now Arctosia arctica). The latter was a new species described by Friele in 1878. Jeffreys thought it to be a mere variety of T. vitrea (T. minor), but Davidson wrote in a letter shortly before he died that Friele's species was distinct (Friele 1886; see Friele 1878, figs 1–2 for comparison.)

The first comprehensive paper on Greenlandic brachiopods was published in 1898, based on an unfinished manuscript by H. J. Posselt. The nec-essary additions and publication were carried out by A. S. Jensen. Posselt's paper is very important, because the author had studied not only the col-lections held in Copenhagen, but also those in the Naturhistoriska Riksmuséet in Stockholm, which were collected by Swedish expeditions. Posselt recorded seven species, six with locali-ties and distribution, mainly based on published

information from British or Norwegian expedi-tions. The seven species were listed as Neatria gnomon, Rhynchonella psittacea, Terebratella spitzbergensis, Terebratulina caput serpentis, Terebratulina arctica, Waldheimia cranium and Terebratulina septentrionalis).

The Greenland brachiopods now held at SNM/ZMUC comprise material collected during the period 1859−98 (not including Chemnitz) by, i.a., the following expeditions: the Konebåd Expedition (1883–85, East Greenland; Holm & Garde 1889), the Fylla Expeditions (1884, 1886, West Greenland; Holm 1889) and the Ingolf Expedition (1895–96, the Davis Strait, South West Greenland, Jan Mayen and the North Atlantic; Wesenberg-Lund 1941, Wolff 2008). Posselt/Jensen commented briefly on the vari-ous species, e.g., R. psittacea (now Hemithiris psittacea) as being the most common species, particularly on hard substrate/bedrock and in slightly deeper water (188 m). T. spitzbergensis (now Glaciarcula spitzbergensis) had only been found very seldomly. Posselt separated T. sep-tentrionalis (American) from T. caput serpentis (now T. retusa) based on shell sculpture. The latter species is said to be very rare.

In 1898–1900, Amdrup led the East Green-land Expedition to the area north of Angmagssa-lik (Amdrup 1902). More than 14 specimens of H. psittacea and about 40 of Liothyris arctica (now Arctosia arctica) were collected by the schooner Godthaab in Forsblad Fjord and south of Shannon Island (Wesenberg-Lund 1940b). A Swedish zoological polar expedition to Spitzber-gen, northeastern Greenland and Jan Mayen also took place in 1900 (Hägg 1904). It recovered three species, R. psittacea (one dead specimen), T. caput serpentis (five specimens) and T. arctica (55 specimens, "dead and alive") from four loca-

Fig. 2. Drawing of Hemithiris psittacea by H. P. C. Möller. SNM/ZMUC archive. Photo by E. Thomsen, TMU.

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tions off East Greenland. Comparing T. caput serpentis (now retusa) and T. septentrionalis, Hägg believed they were varieties of the same species.

In 1908, Blockmann published a paper on the systematic and geographical distribution of Recent brachiopods. This paper included a de-tailed study of Liothyris arctica (now A. arctica) (Blockmann 1908: 613−614), accompanied by a map showing its distribution in East Greenland. A few specimens had already been recorded by Grieg (1907: 540) from a station off East Green-land, based on material from the Belgica Expedi-tion, which took place in 1905.

The Danmark Expedition in 1906−08 to North East Greenland (Amdrup 1913) recovered

five specimens of H. psittacea in Sydostbugten in 1906. Eight complete individuals and some valves of H. psittacea were collected in Storm-bugten in 1907 (Wesenberg-Lund 1940b).

Dall (1920) presented an annotated list of Recent Brachiopoda in the collection of the Smithsonian Institution (National Museum of Natural History). Among these was the mate-rial of seven species from Greenlandic waters collected by, i.a., Jeffreys during the Valorous expedition. Jeffreys' collection had been pur-chased by the museum. Other contributors to the collection were Friele and Mörch. Specimens of Hemithiris psittacea from Valorous dominated the collection. Cooper (1983) used Greenlandic material from the collection of the Smithsonian in his description of a new genus, Arctosia, with Terebratula arctica Friele, 1878 as type species.

A brief note on a likewise small material, a single specimen of Waldheimia cranium (now M. cranium) from the Holsteinsborg area, West Greenland, was published by Schlesch (1925). This note also referred to specimens collected by the Swedish Sofia Expedition in 1871, from the Julianehåb area (60°43' N).

Grieg published a synoptic paper on the Sval-bard brachiopods in 1933. It was primarily based on material collected around Svalbard during the period 1923−31, but also included/listed material from Norwegian expeditions: the Norwegian North Atlantic Expedition (Vöringen off Jan Mayen, 1876−78), the North Atlantic Deep-Sea Expedition (Michael Sars, 1910), and the Veiding Expedition (the Denmark Strait and Jan Mayen, 1931). The material comprised five species from West Greenland and four from East Greenland.

Helmcke presented a paper on the brachiopods of the Zoological Museum in Berlin in 1939, recording a sparse material from Greenland that consisted of 14 specimens in three species: Terebratulina septentrionalis, Macandrevia cra-nium and Hemithiris psittacea (as Hemithyris).

Shortly afterwards, in 1940, M. E. Wesenberg-Lund published two monographic papers on the brachiopods of West and East Greenland respectively. These papers were the first synoptic papers on Greenland brachiopods since Posselt's in 1898. They are the first to list all the material stored in ZMUC, collected by Danish expedi-tions and other collectors since 1842, including

Fig. 3. Hand colored drawing of Hemithiris psittacea by H. P. C. Möller. SNM/ZMUC archive reference B 147E. Photo by E. Thomsen, TMU.

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the material listed by Posselt (1898). Today the museum possesses material dating from (before) 1785 to 1992. The present paper gives the first complete list of this material.

Her paper on the brachiopods of West Green-land (Wesenberg-Lund 1940a), besides older material in the collections of ZMUC, included material from expeditions with Ingolf (1895−96), Dana (1922, 1925, 1935) and Godthaab (1928, 1932−33), as well as from investigations of sev-eral fjords: Tjalfe (A. S. Jensen, 1908–09) and Rink (Nordre Strømfjord near Holsteinsborg, Nordmann 1911; Kvanefjord, Bredefjord and Skovfjord in 1912, Stephensen 1917). Her pa-per, however, was initiated by the investigations of the material collected by Godthaab in 1928. The specific aim of this expedition was to study the waters west of Greenland, in particular the hydrography and biology on both sides of a sub-marine ridge across the Davis Strait between Hol-steinsborg and Cape Walsingham in Baffinland.

Wesenberg-Lund recorded six species as Pelagodiscus atlanticus, Hemithyris psittacea, Neatria gnomon, Terebratulina septentriona-lis, Terebratella spitzbergensis and Waldheimia (M.) cranium. The occurrence of T. retusa was considered dubious because of confusion with T. septentrionalis. This paper provided remarks on morphology, epifauna, substrate and distribution. The publication of the fjord material is of great importance because it demonstrated the occur-rence of two different types of fjords, the "Arctic" type (with very cold, arctic bottom water behind a ridge at the mouth) and the "Atlantic" type (with no ridge and free access for warm Atlantic water). The types were discovered as a result of the investigations of Tjalfe.

Following this paper, Wesenberg-Lund pub-lished a similar synopsis on the brachiopod spe-cies of East Greenland, based on the collections at ZMUC (Wesenberg-Lund 1940b). The mate-rial consisted mainly of brachiopods collected on expeditions during the 1930's, i.e., expeditions that mainly concentrated on studying the animal communities of the coastal waters and fjords from Lindenowfjord (61° N) with Thor in 1935 (Bertelsen in Gjessing et al. 1937), to Danmarks-havn (about 77° N). Five species were recorded as H. psittacea, L. arctica, T. spitzbergensis, T. septentrionalis and M. cranium. Additional

material (previously mentioned) was recorded from the East Greenland Expedition (1898−1900) and the Danmark Expedition (1906−08). T. spitz-bergensis (now Glaciarcula spitzbergensis) was new to East Greenland, but was been recorded as two dead specimens from Lindenowfjord. Hence, Wesenberg-Lund doubted whether this was a true East Greenlandic species.

Since these two papers were published by Wesenberg-Lund, new records of Greenlan-dic brachiopods have been sporadic, e.g., Vibe (1950), Cooper (1973), Marin I.D. (1978) and Piepenburg (1988). The material presently stored at ZMUC, however, also reveals major collec-tions of unpublished, unregistered material from Danish expeditions, e.g., the Thule Expedition in 1968 (Just 1980), the Kap Farvel Expedition in 1970 (NN 1970, Røen 1971) and the Greenland Expedition (Dana) in 1975−78. In addition, the Arctic Biological Field Courses held annually at the station at Godhavn/Qeqertarsuaq, Disko Island, have contributed to the collection (e.g., Petersen 1985).

Vibe participated in the Danish Thule and Ellesmere Land Expedition (1939−41). The purpose of this expedition was to carry out zoo-logical, botanical and geological investigations. Following the outbreak of World War II, marine fieldwork became restricted due to lack of fuel for the small open boat used by the expedition. Nevertheless, shallow water communities were sampled by simply lowering the grab though holes in the ice and getting the dogs to haul it up. This enabled a rich marine sublittoral fauna to be listed and he documented a Macoma cal-carea community, which included the brachio-pod Hemithiris psittacea (1−2 specimens/m2) from bottoms consisting of clay, mud and gravel-sized dropstones (Fig. 4).

In 1968, Vibe and Just participated in the Dan-ish Atomic Energy Commission's Radio-ecologi-cal Expedition to Thule in North West Greenland and collected material in the Thule area (Just 1980). The main sampling area was Bylot Sund. Altogether, 11 stations were sampled (fig. 1 in Just 1980) and specimens of H. psittacea and G. spitzbergensis were recovered, the first being the most frequent. The sampling depth of the brachiopod stations ranged from 22 to 210 m (up to 300 m in Murchison Sound). The brachiopod

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material from this and the following expeditions has not been published.

The Danish Research Council and the De-partment for Greenland supported a biological expedition in 1970 to the Kap Farvel area in the southern part of Greenland (NN 1970, Røen 1971). This is considered a key area for com-parisons of the southern west and east coasts. A small private fishing boat was used. Just, Lemche and Nørrevang collected the largest number of

brachiopods ever from Greenland and further-more, the samples turned out to be important, as they also comprised many juvenile individuals, previously often lost due to sampling methods. The sampling area from Prins Christiansund to Torssukátak is subject to strong currents, result-ing in a high arctic epifauna with a large distri-bution and rich communities of, e.g., hydroids, sponges, bryozoans and brachiopods. Brachio-pods were sampled from 27 stations. Most com-

Fig. 4. Macoma calcarea community from the Thule district. Note two circled specimens of the brachiopod Hemithiris psit-tacea. From Vibe 1950.

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mon was H. psittacea, which occurred together with T. septentrionalis and G. spitzbergensis. Most specimens were recovered from depths ranging from less than 150 m to 200 m.

Cooper (1973) identified 32 species of bra-chiopods from the cruises of the R/V Vema in Smithsonian material. Among these were T. septentrionalis and M. cranium from three locali-ties southwest of Greenland. He also noted that brachiopods sometimes dominate the benthic fauna, particularly in the Antarctic. Zenkevitch (1963) had a similar experience describing a Ma-candrevia community in the eastern Barents Sea.

A later report (Marin I.D. 1978) recorded all benthic invertebrates in material that had been collected during 1975−76 by Dana and Adolf Jensen between 63° N and 68° N in West Green-land. Four brachiopod species were recorded: Novocrania anomala (as Crania, one specimen; record considered dubious because this is a shal-low water species and the depth at the station was 176 m), Hemithiris psittacea, (as Hemithyris) Glaciarcula spitzbergensis (as Diestothyris) and Cryptopora gnomon (one specimen). The most common species was H. psittacea, primarily found in depths between 24 and 74 m. Altogether, 68 specimens of brachiopods were recovered.

The Arctic Station in Godhavn, West Green-land, was founded in 1906 and has since provided logistics for many expeditions and field courses from the University of Copenhagen (Bruun et al. 2006). The marine biology course there in 1984 concentrated on the Disko Bay area, between 69° N and 70° N, and recovered H. psittacea in depths of 30−100 m (Petersen 1985).

Piepenburg (1988) recorded two brachiopod species, H. psittacea and W. cranium (now M. cranium), from a cruise with R/V Polarstern in 1985 off North East Greenland. The brachiopods were in most cases found as empty shells in the Belgica Bank area, between 78° N and 80° N (Thomsen 1990).

Thomsen (1989, 1990) studied the application of brachiopods in palaeoceanographic recon-structions, in particular Macandrevia cranium and Hemithiris psittacea. The biogeography of the former species, including occurrences in Greenlandic water, shows an almost complete confinement to the shelves surrounding the

North Atlantic and the Norwegian-Greenland Sea (Thomsen 1990). This study provided substantial evidence that this distribution is a function of dispersal by the Gulf Stream and its continuations; it is thus connected with Atlantic water masses (temperatures above 7° C), con-firming that brachiopods can be used as markers in palaeoceanographic reconstructions. Curry & Endo (1991) studied the distribution of T. retusa and T. septentrionalis in the North Atlantic. They concluded that T. retusa has its main distribution in the eastern North Atlantic and T. septentriona-lis in the western North Atlantic, including East Greenlandic waters.

MATERIAL AND METHODS

All the material reported on here is housed at the Zoological Museum of the University of Copen-hagen (ZMUC), now part of the Natural History Museum of Denmark (SNM).

The classification used follows the Treatise on Invertebrate Paleontology 6 (H): Brachiopoda 1965 and Brachiopoda revised 1997, 2000, 2002, 2006 and 2007.

No attempt has been made to present a com-plete list of synonymies for the species treated, as this may be found elsewhere, since the species are all well known. However, the original de-scription followed by all published Greenlandic references are listed.

In addition to listing the material examined for each species, information is also provided on material that was expected to be present in the ZMUC collection but not found there.

Legend: ex = complete specimen, ex+ = complete dead specimen, pv = pedicle valve, bv = brachial valve. The geographical division into North (N), West (W) and East (E) follows the official province boundaries (cf. Holmberg & Jensen 2009). A "ca." before a coordinate means that it has been calculated.

TAXONOMY

Class LingulataOrder LingulidaFamily Discinidae

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Pelagodiscus atlanticus (King, 1868)Fig. 5

Discina atlantica King, 1868: 170.Pelagodiscus atlanticus. – Dall 1920: 280. – Wesenberg-

Lund 1940a: 5−6, fig. 1.

Material examined:W GREENLAND (ZMUC): 7 ex, Ingolf 1895 st. 36, 61°50' N, 56°21' W, 2700 m.

DescriptionsDescriptions in King (1868), Jeffreys (1876b), Wesenberg-Lund (1940a) and Brunton & Curry (1979).

RemarksThe species is attached to hard substrates by cementation of the ventral valve. Some of the Greenland specimens were attached to small pieces of basalt (Wesenberg-Lund 1940a).

Class RhynchonellataOrder RhynchonellidaFamily Cryptoporidae

Cryptopora gnomon Jeffreys, 1869Fig. 6

Cryptopora gnomon Jeffreys, 1869: 136.Atretia gnomon. − Jeffreys 1876a: 184, 189, 199−200; 1876b:

251−252. − Dall 1920: 293.Neatretia gnomon. – Posselt 1898: 1−2. − Wesenberg-Lund

1940a: 7−8, fig. 2.

Material examined:W GREENLAND (all ZMUC): 11 ex, Ingolf 1895 st. 36, 61°50' N, 56°21' W, 1665 m; 1 pv, 1 bv, Rink 1911 st. 28, inner parts of Nordre Strømfjord, ca. 67°53' N, 50°14' W, 20−36 m; 1 dubious ex, Godthaab 1928 st. 61, 72°33.5' N, 57°36' W, 190 m; 1 dubious ex, Kap Farvel 1970 st. 40, Kangikitsoq, head of fjord, 60°19.7' N 44°15.7' W, 100 m.

Not present in ZMUC:W GREENLAND: ca. 50 ex, Ingolf 1895 st. 24, 63°06' N, 56°00' W, 2058 m; 1 juvenile ex, Rink 1911 st. 23, Nordre Strømfjord, Ugssuit, ca. 67°48' N, 49°57' W, 50−52 m.

DescriptionsDescriptions in Jeffreys (1876a and b), Wesen-berg-Lund (1940a), and Brunton & Curry (1979).

RemarksCryptopora gnomon lives attached to rocks, polychaete tubes, etc. by a long, threadlike ped-icle. The species is probably not documented representatively in various collections because it is difficult to sample, being a very small bra-chiopod (max 6−7 mm in length) with a shell that easily fragments.

Family Hemithirididae

Hemithiris psittacea (Gmelin, 1791)Figs 1−3, 7, 8A, B

Anomia psittacea Gmelin, 1791: 3348 (type locality: "mari Groenlandiam").

T. psittacea. – Möller 1842: 23.Terebratula psittacea. – Möbius 1874: 253.Rhynchonella psittacea. – Jeffreys 1876a: 183. − Norman

1876: 209. – Posselt 1898: 2−3. – Hägg 1904: 1−2.

Fig. 5. Pelagodiscus atlanticus. From Wesenberg-Lund (1940a). Drawing by Poul H. Winther.

Fig. 6. Cryptopora gnomon. From Wesenberg-Lund (1940a). Drawing by Poul H. Winther.

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Hemithyris psittacea. – Dall 1920: 284−286. – Helmcke 1939: 262−263. − Wesenberg-Lund 1940a: 6−7; 1940b: 4−5. − Vibe 1950: 111, fig. 12. – Piepenburg 1988: 40.

Hemithiris psittacea. – Thomsen 1990: 31; 2001: 115−116.

Material examined:Type materialSyntypes: 2 pv and 2 bv, Davis Strait, W Greenland, Chem-nitz (Spengler) no. 13, collected before 1785, ZMUC-BRA-1 to -4 (Fig. 7).Other material GREENLAND (all ZMUC): 2 ex; 42 ex, 4 pv, 2 bv, Chr. VIII's collection; 4 ex, C. M. Poulsen's collection; 2 ex, Zimmer; 1 ex, Rudolph.W GREENLAND (all ZMUC): 8 ex, Möller 1840 no. 1186, Godthaab, ca. 64°10' N, 51°45' W; 12 ex, Möller no. 1630, Fiskernæs, ca. 63°5'7" N, 50°40'39" W; 3 ex, 1 pv, Holböll, Godthaab, ca. 64°10' N, 51°45' W; 12 ex, Olrik, 1859, Riten-benk, ca. 69°46' N, 51°19' W, 188 m; 7−8 ex, Olrik, 1860, Ritenbenk, ca. 69°46' N, 51°19' W and Prøven, ca. 72°23' N, 55°32' W; 10 ex, Andersen, 1863, Ritenbenk, ca. 69°46' N, 51°19' W; 1 ex, Fylla 1884, st. 5, Davis Strait, 66°32' N, 55°34' W, 188 m; 2 ex, 4 pv, Fylla 1884, st. 11, Jacobshavn, ca. 69°13' N, 51°06' W, 19−28 m; 4 ex, Fylla 1887?, Ikertokfjord, South of Holsteinsborg, ca. 66°46' N, 53°27' W, 56 m; 7 ex, Jørgensen, Julianehaab, ca. 60°43' N, 46°2' W, 75−94 m; 1 ex, Wandel, 1889, Davis Strait; 2 ex, Traustedt, 1892, Pecten-banken, Holsteinsborg, ca. 66°56'22'' N, 53°40'24'' W; 5 ex, Traustedt, 1892, Egedesminde, ca. 68°42'35" N, 52°52'11" W; 27 ex, Traustedt, 1892, Ritenbenk, ca. 69°46' N, 51°19' W; 8 ex, 3 ex+, Traustedt, 1892, Holsteinsborg, ca. 66°56'22"

N, 53°40'24" W; 1 ex, Ingolf 1895, st. 25, 63°30' N, 54°25' W, 1096 m; 1 ex, Ingolf 1895, st. 29, 65°34' N, 54°31' W, 128 m; >45 ex, Ingolf 1895, st. 23-7, outlet of Ameralikfjord, ca. 60°43' N, 56°0'W; 1 ex, Rink 1911, st. 2, outer part of Nordre Strømfjord, ca. 67°36' N, 53°08' W, 345 m; 1 ex, Rink 1911, st. 3A, outer part of Nordre Strømfjord, ca. 67°40' N, 53°00' W, 325−330 m; 6 ex, Rink 1911, st. 26, inner part of Nordre Strømfjord, ca. 67°52' N, 50°09' W, 33−50 m; 7 ex, Rink 1911, st. 26-7, inner part of Nordre Strømfjord, ca. 67°52' N, 50°09' W, 35−46 m; 2 ex+, Rink 1911, st. 27, inner part of Nordre Strømfjord, ca. 67°53' N, 50°09' W, 75 m; 1 ex, 1 ex+, Rink 1911, st. 31, inner part of Nordre Strømfjord, ca. 67°51' N, 50°17' W, 35−46 m; 1 ex, Rink 1911, st. 32, inner part of Nordre Strømfjord, ca. 67°47' N, 50°40' W, 20−35 m; 2 ex, 6 ex+, Rink 1911, st. 33, inner part of Nordre Strømfjord, ca. 67°46' N, 50°42' W, 150 m; 1 ex+, Rink 1911, st. 35, inner part of Nordre Strømfjord, ca. 67°45' N, 51°01' W, 210−225 m; 1 ex, Rink 1912, st. 1, Kvanefjord, ca. 62°03' N, 49°28' W, 84 m; 2 ex, Rink 1912, st. 79, Bredefjord, 60°42' N, 46°29' W, 30−50 m; 1 pv, 1 bv, Rink 1912, st. 103, Bredefjord, 60°51' N, 46°21' W, 90−100 m; 3 ex, Rink 1912, st. 109, Bredefjord, 60°51' N, 46°15' W, 125−140 m; 6ex, 4 ex+, 1 bv, Rink 1912, st. 134, Bredefjord, 60°54' N, 46° 04' W, 85−140 m; 4 ex+, Rink 1912, st. 143, Skovfjord, 60°48' N, 46°06' W, 65−90 m; 1 pv, Dana 1925, st. 2363, 69°30' N, 56°32' W, 202 m; 2 ex, 1 bv, Godthaab 1928, st. 33A, off harbour of Holsteinsborg, ca. 66°56'22" N, 53°40'24" W, 35 m; 3 ex, Godthaab 1928, st. 48, 69°41' N, 55°30' W, 120 m; 1 ex+, Godthaab 1928, st. 60, 72°30' N, 59°02.5' W, 245 m; 6 ex, 4 ex+, 1 pv, 2 bv, Godthaab 1928, st. 188, 60°22' N, 47°27' W, 120 m; 1 ex+, Johansen, 1931, bay outside Tasiusak harbour, north of Upernavik, ca. 60°11'35" N, 44°49'11" W, ca. 28 m; 13 ex, 1 pv, Høpner Petersen, 1959, st. 258, Godhavn, along the inner Udkiggen, ca. 69°14' N, 53°34' W, 50−100 m (Fig. 8B); 3 ex, Høpner Petersen, 1959, st. 354, Godhavn, at Udkiggen, ca. 69°14' N, 53°34' W, 60−90 m; 1 ex, Høpner Petersen, 1960, st. 435, Godhavn, at Udkiggen, ca. 69°14' N, 53°34' W; 1 ex, Kap Farvel 1970, st. 11, Ilua, 60°09.4' N, 44°14.9' W, 60−70 m; 5 ex, 2 ex+, Kap Farvel 1970, st. 23, Ilua, 60°09.4' N, 44°14.9' W, 100−120 m; 5 ex, Kap Farvel 1970, st. 25, Ilua, 60°09.4' N, 44°14.9' W, ca. 150 m; 9 ex, 1 ex+, Kap Farvel 1970, st. 40, Kangikitsoq, head of fjord, 60°19.7' N, 44°15.7' W, 100 m (Fig. 8A1−4); 1 ex, 1 ex+, Kap Farvel 1970, st. 41, Kan-gikitsoq, head of fjord, 60°19.7' N, 44°15.7' W, 100 m; >27 ex, 82 ex+, Kap Farvel 1970, st. 61, south of Augpilagtoq,

Fig. 7. Specimens of Hemithiris psittacea from Greenland. Syntypes (ZMUC-BRA-1 to -4). Photo by E. Thomsen, TMU.

Fig. 8 (opposite page). A1−4. Hemithiris psittacea (ZMUC-BRA-5) from Kangikitsoq, West Greenland. B. Several specimens of H. psittacea (ZMUC-BRA-6) from Udkiggen, Godhavn, West Greenland. C. Several specimens (ZMUC-BRA-7) of Terebratulina septentrionalis and one specimen of Macandrevia cranium from Kangamiut, West Greenland. D1−3. T. septentrionalis (ZMUC-BRA-8) from Imerdlugtoq, West Greenland. E. T. septentrionalis (ZMUC-BRA-9) from Dana st. 7274, West Greenland. Note the juvenile sheltering on the adult. F. Several specimens of M. cranium (ZMUC-BRA-10) from Dana st. 2361, West Greenland. G1−4. M. cra-nium (ZMUC-BRA-11) from Dana st. 2361, West Greenland. H1−4. M. cranium (ZMUC-BRA-12) from Dana st. 2361, West Greenland. Photos by T. A. Wacko, TMU.

173brachiopoda groenlandica

e. thomsen174

60°08.5' N, 44°17.4' W, 50−60 m; 1 ex, Kap Farvel 1970, st. 82, tip of point in Qornoq, 60°07.2' N, 44°17.2' W, 50 m; 21 ex, 11 ex+, Kap Farvel 1970, st. 88, Nûa, 59°53.5' N, 44° 22.3' W, 140 m; 1 ex, Kap Farvel 1970, st. 100, Qagik, 60°10.1' N, 44°13.0' W, 40−60 m; 3 ex, 1 ex+, Kap Farvel 1970, st. 101, Qagik, 60°10.1' N, 44°13.0' W, 40 m; 1 ex, Kap Farvel 1970, st. 114, west point of Sangmissoq, 59°59.3' N, 43°55.9' W, 100 m; 7 ex, Kap Farvel 1970, st. 115, west point of Sangmis-soq, 59°59.3' N, 43°55.9' W, 200 m; 1 ex, Kap Farvel 1970, st. 117, Qeqertat southwest islands, 59°52.8' N, 43°31.5' W, 225 m; 3 ex, Kap Farvel 1970, st. 121, Sorratit, east side of point, 59°54.4' N, 43°34.5' W, 60 m; 3 ex, Kap Farvel 1970, st. 129, Imarssuaq, 60°01.7' N, 43°59.3' W, 530 m; 11 ex, 15 ex+, Kap Farvel 1970, st. 132, Pamiagdluk Kujatingua, 59°55.5' N, 44°23.0' W, 400−450 m; 1 ex, Kap Farvel 1970, st. 136, southernmost island in Toqulineq, 60°04.5' N, 43°02.7' W, 240 m; 6 ex, 5 ex+, Kap Farvel 1970, st. 141, west side of Qagssit Avalequta, 60°01.8' N, 43°05.8' W, 100 m; 4 ex, Kap Farvel 1970, st. 142, north of P. C. weather station, 60°03.8' N, 43°09.4' W, 120 m; 23 ex, Dana 1975, st. 53C, 67°05' N, 54°45' W, 70 m; 7 ex, 2 ex+, Arktisk Biologi Feltkursus 1984, st. 1007, outside Udkiggen, south of Godhavn, ca. 150 m from land, ca. 69°14' N, 53°34' W, 56−72 m; 1 ex, 7 ex+, 5 pv, 6 bv, Arktisk Biologi Feltkursus 1984, st. 1057, Fortune Bay near Skarve Island, at Godhavn, ca. 69°18' N, 53°53' W, 30−40 m; 1 ex, 6 ex+, Arktisk Biologi Feltkursus 1984, st. 1057, Fortune Bay near Skarve Island, at Godhavn, ca. 69°18' N, 53°53' W, 30−40 m; 4 ex, Arktisk Biologi Feltkursus 1984, st. 1060, Fortune Bay, deep groove in middle of bay, ca. 69°18' N, 53°53' W, 55−68 m; 6 ex, Arktisk Biologi Feltkursus 1984, st. 1075, Fortune Bay, between Skarve Island and Oqarsorfik, ca. 69°18' N, 53°53' W, 45−55 m; 2 ex, Arktisk Biologi Feltkursus 1984, st. 2002, Udkiggen, ca. 69°14' N, 53°34' W, 90 m; 5 ex, Arktisk Biologi Feltkursus 1984, st. 2003, off Kød Island, ca. 69°14' N, 53°34' W, 90 m.E GREENLAND (all ZMUC): 1 ex, Konebådsekspeditionen 1884−1885, Suarissok; 1 ex, East Greenland Expedition 1899, Tasiusak (Tasissaq), ca. 65°35' N, 37°32' W; 1 ex, East Greenland Expedition 1899, Tasiusak (Tassisaq), ca. 65°35' N, 37°32' W, 28/38−56 m; 1 ex, East Greenland Expedition 1900, Ryders Land, 565 m; 1 ex, East Greenland Expedition 1900, Henry Land, ca. 69°34' N, 23°35' W or ca. 69°35' N, 23°35' W; 1 ex, 1 pv, East Greenland Expedition 1900, Fors-blad Fjord, ca. 72°28' N, 25°10' W or ca. 72°27' N, 25°23' W, 94−170 m; 7 ex, East Greenland Expedition 1902, Tasiusak (Tasissaq), ca. 65°35' N, 37°32' W, 56−94 m; 1 ex+, East Greenland Expedition 1902, Inginerkertok (Ingimikertoq?), Angmagssalik Fjord, ca. 65°45' N, 36°58' W; 5 ex, Danmark Expedition 1906, Sadnsamiafikk, Sydostbugten; 7 ex, 3 pv, Danmark Expedition 1907, st. 63, Stormbugten, ca. 76°45' N, 19°0'W, 19−38 m; 1 ex, Danmark Expedition 1907, st. 66, Stormbugten, ca. 76°45' N, 19°0' W, 38 m; 7 ex, 7 ex+, Dana 1922, Angmagssalik, outside harbour, ca. 65°36'52" N, 37°38'12" W, 45 m; 12 ex, 1 ex+, 7th Thule Expedition 1932, north of Lindenowfjord, 10−75 m; 1 ex, 1 bv, 7th Thule Expedition 1933, st. 4, Sermilik, 200 m; 1 ex+, 7th Thule Expedition 1933, st. 19, Tasissaq, ca. 65°35' N, 37°32' W, 50−75 m; 1 ex, 7th Thule Expedition 1933, st. 21, Tasissaq, ca. 65°35' N, 37°32' W, 100 m; 4 ex, 1 ex+, Thor 1935, st. 10-d, Kekertatsiak, ca. 60°9' N, 43°1' W, 60−70 m; 10 ex, Thor 1935, st. 10-e, Kekertatsiak, ca. 60°9' N, 43°1' W, 60−70 m;

1 ex, Thor 1935, st. 11-2, Lindenowfjord, ca. 60°31' N, 43°40' W, 60−70 m; 10 ex, Thor 1935, st ?, Kekertatsiak, ca. 60°31' N, 43°40' W, 60−70 m.N GREENLAND (all ZMUC): 3 ex, 2nd Thule Expedition 1917, Wolstenholme Sund, ca. 76°37'55" N, 68°38'15" W, 66 m; 3 ex, 1 pv, Godthaab 1928, st. 97, 78°15.5' N, 73°29' W, 290 m; 1 ex, Godthaab 1928, st. 107, 76°25' N, 69°38' W, 165 m; 6 ex, 1 ex+, Godthaab 1928, st. 114, 76°40' N, 74°20' W, 250 m; 6 ex, 4 ex+, Godthaab 1928, st. 116, 76°08' N, 80°53' W, 80 m; 1 ex, 1 ex+, Danish Thule and Ellesmere Land Expedition 1939, Neqèx (probably Neqe), ca. 77°52' N, 71°37' W, 25−40 m; 10 ex, 3 ex+, Thule 1968, st. 12, Bylot Sund, 76°32.9' N, 69°00.2' W, 40−60 m and 100−110 m; 1 ex+, Thule 1968, st. 14, Bylot Sund, 76°32.9' N, 69°01' W, 210 m; 1 ex, Thule 1968, st. 16, Bylot Sund, 76°30.9' N, 69°28.6' W, 88 m; 5 ex, Thule 1968, st. 20, Bylot Sund, 76°34.5' N, 69°24.5' W, 22−35 m; 9 ex, 1 ex+, Thule 1968, st. 26, between Wolstenholme Island and Saunders Island, 76°30.3' N, 69°53.6' W, 34 m; 1 ex, Thule 1968, st. 27, 76°29.9' N, 69°19.0' W, 30 m; 8 ex, Thule 1968, st. 29, Bylot Sund, 76°28.65' N, 69°21.8' W, 97 m; 1 ex, Thule 1968, st. 34, 76°30.1' N, 69°10.7' W, 125 m; 1 ex, Ingolf 1980, st. 21, north of Kap York, 75°59' N, 67°10' W, 41 m.

Not present in ZMUC:E GREENLAND: 1 ex, Kangerdlugssuaq area, south of Kap Dalton, 150 m; 1 ex, off Kangerdlugssuaq, 50 m; 6 ex, Storm-bugten, ca. 76°45' N, 19°0' W, 20−35 m; 6 ex, Stormbugten, ca. 76°45' N, 19°0' W, 40 m.

Description of syntypes

The syntypes comprise two brachial valves and two pedicle valves from four individuals. They are photographed with a tiny pebble that was in the sample; it may have been a substrate (Fig. 7). Three valves are from adult individuals and one pedicle valve is from a juvenile (Figs 1, 7).

The brachial valves are 2,08 and 1,95−1.98 cm long and 2.21 and 2.15 cm wide, respectively.

The pedicle valves are 1.17−1.2 and 2.08 cm long and 0.91 and 1.69−1.72 cm wide, respec-tively.

Descriptions

Descriptions in Wesenberg-Lund (1940a) and Brunton & Curry (1979).

Remarks

The valves of the largest specimens serve as substrate for a prolific epifauna of bryozoans, foraminiferans, serpulids and cirripedians. Nati-cid gastropod borings, e.g., Oichnus isp., are common.

175brachiopoda groenlandica

Order TerebratulidaFamily Terebratulidae

Arctosia arctica (Friele, 1878)Fig. 9

Terebratula arctica Friele, 1878: 221, fig. 1a–c. – Posselt 1898: 6–7. – Hägg 1904: 5.

Liothyris (Terebratula) arctica. – Grieg 1907: 540.Liothyris arctica. – Blockmann 1908: 613–614, map. – Grieg

1933: 67. − Wesenberg-Lund 1940b: 5.Gryphus arcticus. – Dall 1920: 317.Arctosia arctica. – Cooper 1983: 250. Liothyrella arctica. – Zezina 1997: 72.

Material examined:E GREENLAND (all ZMUC): 5–10 ex, East Greenland Expedition 1892, 74°17' N, 15°20' W, ca. 239 m; ca. 10 ex, East Greenland Expedition 1900, Forsblad Fjord, ca. 72°28' N, 25°10' W or 72°27' N, 25°23' W, 94−169 m; 5−10 ex, East Greenland Expedition 1900, Forsblad Fjord, ca. 72°28' N, 25°10' W or 72°27' N, 25°23' W, 94 m; ca. 20 ex, East Greenland Expedition 1900, Kejser Franz Joseph Fjord area, Forsblad Fjord, ca. 72°28' N, 25°10' W or 72°27' N, 25°23' W, ca. 100 m.

DescriptionsDescriptions in Friele (1878), Blockmann (1908), Wesenberg-Lund (1941) and Cooper (1983).

Remarks Sampling may be biased because this species is so fragile and easily crushed.

Family Cancellothyrididae

Terebratulina septentrionalis (Couthouy, 1838)Fig. 8C−E

Terebratula septentrionalis Couthouy, 1838: 65–66.Terebratulina septentrionalis. − Helmcke 1939: 237. –

Wesenberg-Lund 1940a: 9, 12; 1940b: 6−7. − Cooper 1973: 12. – Curry & Endo 1991: 1101−1102. – Thomsen 2001: 118.

Material examined:W GREENLAND (all ZMUC): 6 ex, Holm, 1884, Davis Strait, 188 m; 3 ex, Holm, Davis Strait, 150 m; 2 ex, Wandel, 1889, Davis Strait, 65°22' N, 54°02' W, 124 m; 4 ex, Wandel, 1889, Davis Strait; 14 ex, Ingolf 1895, st. 25, 63°30' N, 54°25' W, 1094 m; 3 ex, Ingolf 1895, st. 27, 64°54' N, 55°10' W, 739 m; 25 ex, Ingolf 1895, st. 28, 65°14' N, 55°42' W, 790 m; 18 ex, Ingolf 1895, st. 32, 66°35' N, 56°38' W, 598 m; 1 ex, Ingolf 1895, st. 33, 65°16' N, 55°05' W, 682 m; 1 ex, Ingolf 1895, st. 34, 65°17' N, 54°17' W, 103 m; 12 ex, Ingolf 1895, st. 35, 65°16' N, 55°05' W, 682 m; 3 ex, Tjalfe 1908, st. 107, 68°20' N, 54°03' W, 360 m; 4 ex, Tjalfe 1908, st. 199, 68°28' N, 54°47' W, 218−245 m; 6 ex, Tjalfe 1909, st. 337, 64°05' N, 55°20' W, 1100 m; 3 ex, Tjalfe 1909, st. 370, 66°45' N, 56°28' W; 3 ex, Tjalfe 1909, st. 397, 66°42' N, 56°12' W, 244 m; 8 ex, Tjalfe 1909, st. 402, 64°40' N, 56°37' W, 720−775 m; 3 ex, Tjalfe 1909, st. 406, 64°22' N, 56°06' W; 1 ex, Tjalfe 1909, st. 407, 64°14' N, 55°55' W, 539 m; 7 ex, Tjalfe 1909, st. 408, 64°14' N, 53°55' W, 839 m; 1 ex, Tjalfe 1909, st. 437, 63°17' N, 52°02' W, 700−1055 m; 2 ex, Rink 1912, st. 45, Bredefjord, 430−450 m; 2 ex, Rink 1912, st. 67, Bredef-jord, 60°38' N, 46°36' W, 220−310 m; 4 ex, Rink 1912, st. 70, Bredefjord, 60°38' N, 46°45' W, 225−290 m; 8 ex, Rink 1912, st. 76, Bredefjord, 60°44' N, 46°36' W, 260−320 m; 1 ex, Rink 1912, st. 77, Bredefjord, ca. 60°41' N, 46°33' W,

Fig. 9. Arctosia arctica (ZMUC-BRA-13) from 74°17'N, 15°20'W, East Greenland. Photos by M. Karlstad, TMU.

e. thomsen176

245−470 m; 2 ex, Rink 1912, st. 91, Bredefjord, 60°48' N, 46°28' W, 110−180 m; 5 ex, Rink 1912, st. 97, Bredefjord, 60°49' N, 46°18' W, 250−280 m; 1 ex, Rink 1912, st. 115, Sermilikfjord, 61°17' N, 45°45' W, 500 m; 1 ex, Rink 1912, st. 135, Bredefjord, 60°54' N, 46°04' W, 225−250 m; 1 ex, Rink 1912, st. 138, Skovfjord, 60°48' N, 45°57' W, 300−360 m; 10 ex, 2 ex+, Dana 1925, st. 2346, 66°37' N, 56°37' W, 450 m; 25 ex, Dana 1925, st. 2361, 68°08' N, 57°30' W, 398 m; 2 ex, Dana 1925, st. 2363, 69°30' N, 56°32' W, 202 m; 17 ex, Godthaab 1928, st. 159, 68°23.5' N, 56°12' W, 480 m; 12 ex, Godthaab 1928, st. 160, 68°17' N, 58°14' W, 410 m; 1 ex, Godthaab 1928, st. 179, 63°36' N, 55°15' W, 1200 m; 3 ex, 1 ex+, Dana 1950, st. 7274, 66°37' N, 57°05' W, 450−493 m (Fig. 8E); 3 ex, Kap Farvel 1970, st. 23, Ilua, 60°09.4' N, 44°14.9' W, 100−120 m; 4 ex, Kap Farvel 1970, st. 25, Ilua, 60°09.4' N, 44°14.9' W, 150 m; 4 ex, Kap Farvel 1970, st. 49, Kangikitsoq, near mouth of fjord, 60°14.9' N, 44°11.6' W, 250 m; 1 ex, Kap Farvel 1970, st. 51, Imerdlugtoq, 60°10.7' N, 44°11.0' W, 300 m (Fig. 8D1−3); 5 ex, Kap Farvel 1970, st. 52, Imerdlugtoq, 60°10.7' N, 44°11.0' W, 200−250 m; 4 ex, Kap Farvel 1970, st. 61, south of Augpilatoq, 60°08.5' N, 44°17.4' W, 50−60 m; 9 ex, Kap Farvel 1970, st. 89, west of Akuiligkat, 59°54.0' N, 44°28.0' W, ca. 400 m; ca 23 ex, 1 ex+, Kap Farvel 1970, st. 128, Imarssuaq, 60°01.7' N, 43°59.3' W, 530 m; 2 ex, Kap Farvel 1970, st. 129, Imarssuaq, 60°01.7' N, 43°59.3' W, 530 m; > 62 ex, Kap Farvel 1970, st. 132, Pamiagdluk Kujatingua, 59°55.5' N, 44°23.0' W, 400−450 m; 4 ex, Kap Farvel 1970, st. 136, southernmost island in Toqulineq, 60°04.5' N, 43°02.7' W, 240 m; 2 ex, OH. 1971, st. ly 711-71-41, 63°32.1' N, 52°59.9' W, 543 m; 8 ex, 1 ex+, Total 1976, Kangamiut 1, x, populated place Kangaamiut, ca. 65°49'30" N, 53°20'33" W (Fig. 8C); 1 ex, Shinkai Maru 1991−92, st. 44, 63°10' N, 54°14' W and 63°10' N, 54°10' W, 1319−1329 m; 4 ex, Shinkai Maru 1991−92, st. 1863, 65°12' N, 56°21' W and 65°10' N, 56°22' W, 732−733 m.E GREENLAND (all ZMUC): 1 ex, 1930, Lichtenaufjord, Nugarsuk, 72°25' N, 17°56' W, 300 m; 1 ex, East Green-land Expedition?, st. 135, Scoresby Sund, outside mouth of Rosenvinges Bugt, 300 m.

DescriptionsDescriptions in Couthouy (1838), Wesenberg-Lund (1941) and Brunton & Curry (1979).

RemarksDall (1920) noted that this species "could gener-ally be distinguished from T. retusa by its finer sculpture and more oval form, but varieties ap-proach one another very closely so far as external aspects is concerned. The usual mutations of form occur very often." Wesenberg-Lund (1940a) confirmed that the two species can be difficult to tell apart morphologically, and that mistaken identity probably can be found in older literature, e.g., Posselt (1898). Later, Wesenberg-Lund (1941) discussed a possible hybridisation of the

morphospecies where their occurrences overlap. She even suggested a name for the hybrid: T. retusa x T. septentrionalis. Based on detailed morphometric studies, however, Curry & Endo (1991) found that T. retusa from Europe and T. septentrionalis from America are two separate species − as confirmed in molecular studies by Cohen et al. (1991) and Lüter & Cohen (2002).

EpifaunaAccording to Dall (1920), this species may serve as a substrate for a ?commensal poriferan which may reduce the adult individuals to "a spongy ball." Tendal & Thomsen (2005) found that a poriferan, Hymesdesmia sp., occurred in the grooves between the radial ribs of primarily adult T. retusa (from the Faroe Islands) suggest-ing a host preference for a ribbed endopunctate brachiopod. Even heavy overgrowth seems nor-mally not to be of inconvenience to the brachio-pod, neither in its movement, nor in its growth and reproduction. A similar poriferan overgrowth is seen in the present material from Greenland, especially of larger specimens.

Terebratulina sp.

Material examined:W GREENLAND (all ZMUC): 1 ex, Kap Farvel 1970, st. 23, Ilua, 60°09.4' N, 44°14.9' W, 100−120 m; 16 ex, Kap Farvel 1970, st. 25, Ilua, 60°09.4' N, 44°14.9' W, ca. 150 m; 2 ex, Kap Farvel 1970, st. 51, Imerdlugtoq, 60°10.7' N, 44°11.0' W, 300 m; 6 ex, Kap Farvel 1970, st. 61, south of Augpilagtoq, 60°08.5' N, 44°17.4' W, 50−60 m; 3 ex, Kap Farvel 1970, st. 102, Akuliaruseq, southeast of point, 60°08.9' N, 44°10.0' W, 250−400 m; 1 ex, Kap Farvel 1970, st. 103, south of Akuliaruseq, 60°08.6' N, 44°10.7' W, 250−400 m; 1 ex, Kap Farvel 1970, st. 115, 59°59.2' N, 43°55.9' W, 200 m; 1 ex, Kap Farvel 1970, st. 129, 60°01.7' N, 43°59.3' W, 530 m; 1 ex, Kap Farvel 1970, st. 136, southernmost island in Toqulineq, 60°04.5' N, 43°02.7' W, 240 m.

Family Dallinidae

Glaciarcula spitzbergensis (Davidson, 1852)Fig. 10A, B

Terebratella Spitzbergensis Davidson, 1852: 78. – Jeffreys 1876a: 182.

Terebratella spitzbergensis. – Posselt 1898: 3−4. – Grieg 1933: 70. – Wesenberg-Lund 1940a: 12, 14; 1940b: 5−6.

Terebratalia spitzbergensis. – Dall 1920: 347.Glaciarcula spitzbergensis. – Thomsen 2001: 120−121.

177brachiopoda groenlandica

Material examined:W GREENLAND (all ZMUC): 2 ex, Godthaab, ca. 64°11'27" N, 51°41'46" W, 56−94 m; 2 ex, Holböll, Egedesminde, ca. 68°42'35" N, 52°52'11" W (Fig. 10A); 1 ex, Møller, Fiske-1 ex, Møller, Fiske-næsset, 63°51'07" N, 50°40'39" W, 103 m; 1 bv, Ingolf 1895, st. 36, 61°50' N, 56°21' W, 2700 m; 2 ex, Rink 1911, st. 31, mouth of Ugssuit, inner part of Nordre Strømfjord, ca. 67°51' N, 50°17' W, 35−46 m (Fig. 10B); 18 ex, Rink 1911, st. 33, 5 miles WSW of Safarssuak, inner part of Nordre Strømfjord, ca. 67°46' N, 50°42' W, 150 m; 1 ex, Rink 1911, st. 35, bay at Spinatarssuak, middle part of Nordre Strømfjord, ca. 67°45' N, 51°01' W, 35 m; 1 ex+, Kap Farvel 1970, st. 2, Qagssialik, 59°54.8' N, 44°26.2' W, 100−120 m; 1 ex, Kap Farvel 1970, st. 14, east of Umiagssat qeqertai, 60°07.8' N, 44°18.4' W, 70−90 m; 1 ex, 1 ex+, Kap Farvel 1970, st. 25, Ilua, 60°09.4' N, 44°14.9' W, ca. 150 m; 1 ex, Kap Farvel 1970, st. 41, Kangikitsoq, head of fjord, 60°19.7' N, 44°15.7' W, 100 m; 10 ex, 9 ex+, Kap Farvel 1970, st. 61, south of Augpilagtoq, 60°08.5' N, 44°17.4' W, 50−60 m; 10 ex, Kap Farvel 1970, st. 88, Nûa, 59°53.5' N, 44°22.3' W, 140 m; 4 ex, Kap Farvel 1970, st. 129, Imarssuaq, 60°01.7' N, 43°59.3' W, 530 m; 1 ex, Kap Farvel 1970, st. 132, Pamiagdluk Kujatingua, 59°55.5' N, 44°23.0' W, 400−450 m; 4 ex+, Kap Farvel 1970, st. 136, southernmost island in Toqulineq, 60°04.5' N, 43°02.7' W, 240 m.E GREENLAND (all ZMUC): 2 ex, Thor 1935, st. 16-4, Lindenowfjord, ca. 60°31' N, 43°40' W, 125−150 m; 1 ex, Lynch, 1972, 67°13.9' N, 18°42.0' W, 221 m.N GREENLAND (all ZMUC): 3 ex, Thule 1968, st. 19, Bylot Sund, 76°34.0' N, 69°19.1' W, 45 m; 1 ex, Thule 1968, st. 68-II, Murchison Sund, 77°23.5' N, 69°29' W, 300 m.

DescriptionsDescriptions in Davidson (1852) and Wesenberg-Lund (1941).

Macandrevia cranium (Müller, 1776)Fig. 8C, F−H

Terebratula cranium Müller, 1776: 249. – Jeffreys 1863: 9, 11−13. – Möbius 1874: 253.

Waldheimia cranium. – Posselt 1898: 7−8. – Schlesch 1925: 148. − Piepenburg 1988: 40.

Macandrevia cranium. – Dall 1920: 354−355. − Helmcke 1939: 250. − Cooper 1973: 24. – Thomsen 1990: 25−37.

Waldheimia (Macandrevia) cranium. – Wesenberg-Lund 1940a: 14; 1940b: 7.

Material examined:W GREENLAND (all ZMUC): 1 ex, Ingolf 1895, st. 27, 64°54' N, 55°10' W, 740 m; 3 ex, Ingolf 1895, st. 34, 65°17' N, 54°17' W, 103 m; 1 ex, Ingolf 1895, st. 35, 65°16' N, 55°05' W, 682 m; 2 ex, Ameralikfjord, ca. 64°07'12" N, 51°0'0" W, 130 m; 1 ex, Rink 1912, st. 67, Bredefjord, 60°38' N, 46°36' W, 220−310 m; 2 ex, Rink 1912, st. 97, Bredefjord, 60°49' N, 46°18' W, 180−250 m; 78 ex, Dana 1925, st. 2361, 68°08' N, 57°30' W, 398 m (Fig. 8F); 1 ex, OSG, st. ly 711-71, no. 41, Northwest Atlantic Ocean, 63°32'106" N, 52°59'54" W, 543 m; 1 ex, Total, 1976, Kangamiut 1, x, populated place Kangaamiut, ca. 65°49'30" N, 53°20'33" W (Fig. 8C).E GREENLAND (all ZMUC): 1 ex, Wandel, Denmark Strait; 2 ex, Wandel, 1889, Denmark Strait, 64°42' N, 27°43' W, 800 m; 10 ex, Ingolf 1895, st. 11, 64°34' N, 31°12' W, 2448 m; 1 ex, Ingolf 1895, st. 91, 64°44' N, 31°00' W, 2350 m; 2 ex, Ingolf 1896, st. 94, 64°56' N, 36°19' W, 384 m; 1 ex, East Greenland Expedition 1900, off Angmagssalik, ca. 65°30' N, 37°20' W, 100 m or 140 m; ca. 100 ex, East Greenland Expedition 1900, Forsblad Fjord, ca. 72°28' N, 25°10' W or 72°27' N, 25°23' W; 1 ex, southeast of Sabine Island, ca. 74°36'0" N, 18°58'59" W, 205 m; 1 ex, Scoresby Sund, Hurry Inlet, ca. 70°39'0'' N, 22°31'59" W, 88 m; 3 ex, 7th Thule Expedition 1932, Kang-erdlugssuak, ca. 68°10' N, 31°45' W, 45 m.

Fig. 10. A. Glaciarcula spitzbergensis (ZMUC-BRA-14) from Egedesminde, West Greenland. B. Glaciarcula spitzbergensis (ZMUC-BRA-15) from the inner part of Nordre Strømfjord, West Greenland. Photos by June Åsheim and M. Karlstad, TMU.

A B

e. thomsen178

N GREENLAND (ZMUC): 1 ex, Ingolf, 74°17' N, 15°20' W, 240 m (no year or station given).

Not present in ZMUC:E GREENLAND: 100 ex, Kejser Franz Joseph Fjord area, 35 m.

DescriptionsDescriptions in Jeffreys (1863) and Brunton & Curry (1979).

RemarksTendal & Thomsen (2005) demonstrated a host preference for epibionts in North Atlantic bra-chiopods. It seems that bryozoans, cirripedians and serpulids all prefer a non-ribbed shell, such

as that of Macandrevia cranium, wheras porif-erans prefer a ribbed shell morphology such as that of Terebratulina retusa, or in the present study, T. septentrionalis. Brachiopods are in fact important, much used biogenic habitats.

CONCLUSIONSeven species are recognized in the brachiopod collection from about 150 localities from Green-land at ZMUC: Pelagodiscus atlanticus (King, 1868) – very sparse, Cryptopora gnomon Jef-freys, 1869 – very sparse, Hemithiris psittacea (Gmelin, 1791) – very abundant, Arctosia arc-tica (Friele, 1878) – very sparse, Terebratulina

Fig. 11. Map showing the distribution of Hemithiris psittacea in Greenland waters. Localities shown in grey refer to localities with calculated coordinates.

179brachiopoda groenlandica

septentrionalis (Couthouy, 1838) – abundant, Glaciarcula spitzbergensis (Davidson, 1852) – sparse, and Macandrevia cranium (Müller, 1776) – sparse. No new species have been found, as the record of Crania anomala (Müller, 1776) (now Novocrania anomala) is considered dubious.

P. atlanticus is listed one time in the record of examined material, C. gnomon four times, H. psittacea 109 times, A. arctica four times, T. sep-tentrionalis 55 times, G. spitzbergensis 20 times and M. cranium 20 times. A few terebratulinids, Terebratulina sp., cannot be recorded at the spe-cies level without further investigations using DNA-methods; they occur nine times in the list.

All the brachiopods from Greenlandic waters

are sessile epibenthic suspension feeders. One species, P. atlanticus, is cemented to its substrate, while all the others are attached by a pedicle to different firm substrates, including conspecif-ics. Thomsen (2005) studied the brachiopod-substrate relationships in the Faroe Islands area.

In order to understand the biogeography of the Greenlandic brachiopods, their geographic distributions were mapped and compared with bathymetric distribution (depth) and temperature of water mass.

Geographic distributionMost of the brachiopods were collected from rather shallow water along the coast, in the fjords

Fig. 12. Map showing the distribution of Terebratulina septentrionalis in Greenland waters. The locality shown in grey refers to a locality with calculated coordinates.

e. thomsen180

or at the ridge between Greenland and Iceland. H. psittacea (Fig. 11) is the only species found all along the coast of Greenland, including some of the fjords, e.g., Nordre Strømfjord and Brede-fjord. T. septentrionalis (Fig. 12) occurs mainly along the west coast south of 70° N, including several fjords, e.g., Bredefjord and Skovfjord. Off the east coast it is only documented by one single specimen from the Scoresby Sund area and one from off Lichtenaufjord further north. G. spitzbergensis (Fig. 13) is found along the west coast from Thule in the north to Kap Farvel in the south, including Nordre Strømfjord. It has also been recorded from off Iceland, but not off

the east coast of Greenland. Wesenberg-Lund (1940b) doubted whether it was a true East Greenland species. A. arctica is, as documented by Wesenberg-Lund (1940b), restricted to the Denmark Strait and the east coast from ca. 67° N to ca. 75° N (Fig. 13). Most of the specimens in the collection are from Forsblad Fjord. M. cranium is primarily known from the west coast south of 70° N and some fjords, e.g., Bredefjord, Skovfjord and Sermilik, but it has also been recorded on the East Greenland shelf (Thomsen 1989, 1990, Fig. 13).

P. atlanticus is only known from one locality off the shelf of West Greenland (Fig. 13).

Fig. 13. Map showing the distribution of five of the brachiopod species that occur in Greenlandic waters. Pelagodiscus atlanticus = square; Cryptopora gnomon = triangle; Arctosia arctica = dot; Glaciarcula spitzbergensis = star and Macandrevia cranium = diamond. Localities shown in grey refer to localities with calculated coordinates.

181brachiopoda groenlandica

C. gnomon is found on the shelf of West Greenland, but also off this shelf in deep water (Fig. 13). The sparse material of P. atlanticus and C. gnomon is not sufficient for a conclusion on the distribution of these species in Greenlandic water.

Bathymetric distribution

There are two species that occur in deep water off the Greenland shelf, P. atlanticus and C. gnomon; the latter, however, has also been recorded from depths less than 200 m on the shelf and from 20−52 m in Nordre Strømfjord. P. atlanticus has a cosmopolitan distribution, occurring in all oceans, always in deep water (366−5482 m according to Brunton & Curry 1979). The only Greenlandic record is from 2700 m. C. gnomon has a North Atlantic distribution, occurring as far south as Cuba (Dall 1920, Brunton & Curry 1979). Wesenberg-Lund (1940a) considered it to be an abyssal form in spite of the known shallow occurrences less than 200 m. These occurrences may be explained by a preference for low tem-peratures. According to Brunton & Curry (1979), the species has a bathymetric range of 183−4023 m. The Greenlandic range is 20−2058 m. H. psit-tacea is a circumpolar species mainly occurring in shallow water on the shelf or in some of the fjords (ca. 80 % from less than 200 m); it has, however, been collected down to 1096 m. It is frequently found together with T. septentrionalis and/or G. spitzbergensis. The bathymetric range for T. septentrionalis is 50−1329 m (ca. 80 % from less than 600 m) and for G. spitzbergensis 35−2700 m (with a preference for less than 300 m). A. arctica occurs at 94−169 m in Forsblad Fjord in East Greenland. The sampling of this species may, however, have been biased and its occurrence/distribution hence difficult to assess. M. cranium has a bathymetric range in Greenlan-dic waters of 35−2448 m (ca. 60 % from less than 300 m). A conclusion must be that Greenlandic brachiopods generally live in depths less than 300 m.

Temperature, current and water massThe investigated area of Greenland is influenced by some major currents, primarily the East

Greenland Current, the Irminger Current and the West Greenland Current. The East Greenland Current protrudes from the Arctic Ocean through the Fram Strait and proceeds along the eastern coast, bringing cold (temperature 0 °C, 3−5 °C in the summer) Polar Water southwards on the shelf to Kap Farvel, where it turns along the west coast northwards to the area between Godthåb/Nuuk and Sukkertoppen/Maniitsoq (Born & Böcher 1999). The Irminger Current carries warm At-lantic Water (6−8 °C) from the North Atlantic Current mixed with local water, the so-called Irminger Water, to the southern shelf of Green-land. This water follows the East Greenland Current, partly below and partly on the outside of the Polar Water, and continues around Kap Farvel into the Davis Strait. The two water masses are mixed along their course, resulting in the rather temperate state of the West Greenland Current that runs northwards along the west coast. The northernmost investigated area, e.g., the Thule/Qaanaaq area, is influenced by Polar Water from the Arctic Ocean protruding southwards through the Nares Strait. This current meets the deflected West Greenland Current and proceeds as the Lab-rador Current along the Canadian coast.

The distribution of Greenlandic brachiopods is closely related to the temperature of the water masses. Thus, A. arctica occurs in areas dominated by the cold Polar Water from the East Greenland Current, i.e., along the east coast and in the Denmark Strait. While occurrences of M. cranium on the southern Greenland shelf may be related to the Irminger Water. The occurrence of the species north of the Greenland-Iceland Ridge on the East Greenland shelf may relate to the influx of Atlantic Water from a branch of the West Spitsbergen Current during the Holocene climatic optimum (Thomsen 1989, 1990). T. sep-tentrionalis occurs in areas influenced by warm Irminger Water; its distribution is thus restricted to the southern part (south of 70°N) of the west coast, including several "Atlantic" fjords, e.g., Bredefjord and Skovfjord, as seen on Fig. 12. H. psittacea is a species with preference for low temperatures (cf. Thomsen 1989, 1990). It is of-ten found together with T. septentrionalis and/or G. spitzbergensis, the latter also being a cold-water species. H. psittacea and G. spitzbergensis

e. thomsen182

are both found in the northernmost West Green-land and in the "Arctic" fjord, Nordre Strømfjord.

The relationship between temperature, cur-rent, water mass and benthic fauna in Greenlan-dic fjords is, as mentioned above, seen in the two types of fjords, originally discovered by the Tjal-fe expedition. The so-called "Arctic" fjords, e.g., Nordre Strømfjord, have bottom temperatures ranging from -1.2 to +0.7 °C, because a ridge prohibits warm “Atlantic Water” from entering the fjord. Kvanefjord, Bredefjord, Sermilikfjord and Skovfjord are typical "Atlantic" fjords with no ridges and bottom temperatures ranging from +0.1 to +3.9 °C (i.e., they are in fact "Arctic-Atlantic" fjords). These types are reflected in the occurrence of the different brachiopod species; thus, M. cranium has only been recorded from the "Atlantic" fjords and G. spitzbergensis has, with one exception, only been found in the "Arc-tic" fjords. H. psittacea occurs in both types of fjords, because it is in fact a species that prefers low temperatures (cf. Thomsen 1989, 1990). In East Greenland, H. psittacea is thus restricted to shallow waters in the "Arctic" fjords.

Key to the Brachiopoda known from Green-land by J. Hansen (pers.comm. 2012)This key includes the seven species which have been reported in this paper.

1 Shell attached to hard substrate by cementa-tion. Shell subconical, inarticulate (no hinge) and with holoperipheral growth. Small, up to 6 mm long, brownish colour, with long setae radiating out from the mantle along the valve margin ......................................................... Pelagodiscus atlanticus (King,1868)

– Shell articulated. Species with pedicle (stalk) ..................................................................... 2

2 Shell impunctate (not punctate). Deltidial plates not meeting. No ornamentation or with striae rather than ribs ................................... 3

– Shell endopunctate (fine pitting seen on shell). No ornamentation or with ribs ..................... 4

3 Uniplicate anterior valve margin. Moderate size (up to 29 mm long). Bluish-brown colour. Fine radiating striae ................................................ Hemithiris psittacea (Gmelin,1791)

– Slightly unisulcate anterior valve margin. Small (up to 7 mm long) and transparent. No ornamentation ............................................................... Cryptopora gnomon Jeffreys, 1869

4 Radiating ornamentation. 15–22 fine ribs per 5 mm 10 mm anterior of ventral umbo. Generally egg-shaped outline .......................................... .. Terebratulina septentrionalis (Couth.,1838)

– Lacking ornamentation. Anterior margin rec- Lacking ornamentation. Anterior margin rec-timarginate. Longer than wide ..................... 5

5 Dorsal valve with median septum. Umbo beak-shaped. Pedicle has rootlets. Up to 15 mm long .. Glaciarcula spitzbergensis (Davidson,1852)

– Median septum lacking in dorsal valve in larg- Median septum lacking in dorsal valve in larg-er individuals. Blunted umbo ...................... 6

6 Up to 31 mm long. Not glassy semitranspar-ent shell. Long brachial loop, supported by septum in small individuals ....................... ............. Macandrevia cranium (Müller,1776)

– Up to 15 mm long. Glassy semitransparent shell. Short brachial loop ........................................................ Arctosia arctica (Friele, 1878)

ACKNOWLEDGEMENTS

The author is grateful to Ole Secher Tendal and Tom Schiøtte (SNM/ZMUC) for continual sup-port and comments on the present paper. Jesper Hansen (AkvaplanNiva-Tromsø) is thanked for the key to the brachiopod species presented in this paper. As former curator of brachiopods, Claus Nielsen kindly provided access to the brachiopod collection at ZMUC. Help by Tom Schiøtte with the maps (Figs 11−13) and Antonia Vedelsby with regard to work in the collection was very much appreciated. Hanne Espersen (li-brarian at SNM/ZMUC) is thanked for help in the library at the museum, which contains a rare col-lection of old and important books, e.g., Chem-nitz (1785). Thomas A. Wacho, June Åsheim and Mari Karlstad (all TMU) and the author took the photos and Ernst Høgtun (TMU) helped with Figs 8 and 10. This study was mainly carried out at SNM/ZMUC, which kindly provided fa-cilities for study. The University of Tromsø, the Research Council of Norway, and private means financed this study.

183brachiopoda groenlandica

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Appendix

Localities: Names of localities cited in this paper are listed in Danish (various spellings)/English/Greenlandic (various spellings), verified when possible using Gaarn Bak (1978) and Jakobsen et al. (2000). Some localities have names that could not be located on geographical maps, e.g., Suarissok; these are indicated by a question mark.

AkuiligkatAkuliarusek/Akuliaruseq/TasiilaqAngmagssalik/AmmassalikAngmagssalik Fjord/Ammassalik FjordBredefjord/Brede Fjord/Nordre Sermilik/

Ikersuak/IkersuaqBylot Sund/Bylot SoundCarlsberg Fjord/CarlsbergfjordDanmarksstræde/Danmarks Stræde/

Denmark StraitDavisstræde/Davis Stræde/Davis StraitDisko/Disko Island/Kekertarssuak/

QeqertarsuaqDisko Bugt/Disko Bay/Qeqertarssûp tunua/

Qeqertarsuup TunuaEgedesminde/Ausiait/AasiaatFiskenæsset/Kekertarssuatsiait/

Qeqertarssuatsiait/QeqertarsuatsiaatForsblad FjordFortune Bay/Fortunebay/KilliitGodhavn/Kekertarsuak/QeqertarsuaqGodthåb/Godthaab/Nûk/NuukHenry LandHolsteinsborg/SisimiutHurry InletIluaIkertokfjord/Ikertôq/Iqertoq FjordImarssuaqImerdlugtoqInginerkertok/Ingmikertok?Jackson Ø/Jackson IslandJakobshavn/Jacobshavn/IllulissatJulianehåb/Julianehaab/Kakortok/QaqortoqKangamiut/KangaamiutKangerdlugssuak (E Greenland) KangikitsoqKap Farvel/UummannarsuaqKejser Franz Joseph FjordKekertat/Qeqertat

KekertatsiakKong Oscars Havn/TasissaqKôrnok/Qôrnok/QoornoqKvanefjord/Kvannefjord/Kuanersok/

Kuanersoq/Kuanersôq/KuannersooqKødø/Kød Ø/QeqertaqLichtenaufjordLindenow Fjord/Lindenowfjord/

Kangerdlugsuatsiak/KangerlussuatsiaqLysefjord/Ameralik Fjord/AmeralikMurchison Sund/Murchison SoundNanortalik/Augpilagtoq/AupilatoqNeqèx? (probably Neqe)Nordre Strømfjord/NassuttooqNûa/NuuaNugarsuk/Nûgârssuk/NugaarsukOqarsorfikPamiagdluk KujatinguaPrins Christian Sund/Prins Christiansund/

Prince Christian Sound/Ikerasarsuak/ Ikerasagssuaq/Ikerasassuaq

Prøven/Kangersuatsiak/KangersuatsiaqQagik?Qagssialik/QassialikQagssit AvalequtaRitenbenk/AppatRyders LandSabine Ø/Sabine IslandSadnsamiafikk?SafarssuakSangmissoq/SammisoqSaunders Ø/Saunders IslandScoresbysund/Igdlorkortôrmiut/

IttoqqortoormiitSermilikfjorden/SermilikShannon Ø/Shannon IslandSkarveø/Skarve Ø/Skarve IslandSkovfjord/Tunulliarfik/NardlunakSorratit?SpinatarssuakStormbugtenSuarissok?Søndre Strømfjord/Kangerdlugssuak/

Kangerdlugssuaq/KangerlussuaqSydostbugten?Tasiusak/Tasiusaq (W Greenland)Thule/Qânâq/Qaanaaq

e. thomsen186

TorssukátakToqulineqUdkiggen/Qáqaliaq/QaqqaliaqUgssuit/UssuitUmiagssat qeqertai/Umaissat QeqertaiUpernivik/UpernavikWolstenholme Ø/Wolstenholme IslandWolstenholme Sund/Wolstenholme Sound