Brachiopoda of Greece: an annotated checklist

Abstract Background Until today, only scattered species records of Brachiopoda from Greece have been included in publications on the Mediterranean brachiopod fauna. These records were mostly based on material collected during marine expeditions in the eastern Mediterranean decades ago, while few recent additional records appear in ecological studies. The aim of this paper was to give the first checklist of brachiopod species of Greece, in the framework of the Greek Taxon Information System (GTIS) initiative of the LifeWatchGreece Research Infrastructure (ESFRI), by reviewing the existing literature. New information Twelve brachiopod species have been found in Greek waters so far. The nomenclature, distribution, fossil records, ecology, and literature sources are discussed for each species.


Introduction
Until Logan's revision of the Mediterranean Brachiopoda (Logan 1979), several records of their extant taxa from the eastern Mediterranean were scattered in old publications on molluscs and in general benthic studies (see Logan et al. 2002 for a detailed publication list). Brachiopod records from Greece were mostly based on material that was collected during the oceanographic expeditions of the French research vessels "Calypso" and "Jean Charcot" in the 1950's and 1960's (Logan 1979, Logan et al. 2002. Since then, few ecological studies on the sciaphilic hard substrate communities reported on Brachiopoda from Greek waters (e.g. Antoniadou and Chintiroglou 2005, Taviani et al. 2011, Gerovasileiou et al. 2015. The aim of the present study was to give an updated, annotated list of Brachiopoda of the Greek seas.

Materials and methods
The checklist of Brachiopoda of Greece (Suppl. material 1) was created in the framework of the Greek Taxon Information System (GTIS) initiative of the LifeWatchGreece Research e-Infrastructure (Bailly et al. 2016, this special collection). Its construction was initially based on the species lists of Mediterranean Brachiopoda compiled by Logan (1979) and Logan et al. (2002). All records in these lists were cross-checked against the World Register of Marine Species (WoRMS Editorial Board 2016) and the Brachiopoda Database (Emig et al. 2016) in order to update taxonomy and nomenclature. Records and information published since 2002 were incorporated in the list. The list is annotated with information regarding species distribution in the Greek seas (i.e. North Aegean, South Aegean, Levantine Sea, Ionian Sea), ecological remarks (i.e. depth and habitat), and the relevant literature sources. Information about fossil records of Brachiopoda from Greece was extracted from Logan et al. (2004), Nielsen et al. (2006, Koskeridou (2007), and references therein. The classification follows WoRMS and Emig et al. (2013).

Class Craniata
Order Craniida

Nomenclature:
Doubtful species, may be a synonym of P. anomioides. See discussion.

Discussion
The checklist of Brachiopoda of Greece comprises 12 species classified into 9 genera, 6 families, 2 orders, and 2 classes. Novocrania turbinata and Platidia davidsoni have been included in the list as doubtful species. Logan and Long (2001) separated the species N. turbinata and N. anomala, which had been synonymized for a long period. Emig (2014) made a synthesis on the issue and, on the basis of molecular data analyzed by Cohen et al. (2014), he concluded that the two species were synonyms. However, he mentions in his discussion that the lack of comprehensive study on morphological characters hinders the conclusion, especially concerning the eastern Mediterranean basin. The synonymy of P. davidsoni with P. anomioides has been also questioned (A. Logan, pers. comm.). Thus, we chose to keep the two species in the list until their status is completely clarified.
The only recent addition to the brachiopod diversity of Greece is the species Gwynia capsula (Antoniadou and Chintiroglou 2005). Logan et al. (2004) had attributed the fact that this species was not found in the eastern basin either to the eastward Mediterranean faunal impoverishment or to the limited research effort in this area on this phylum. Indeed, G. capsula was reported one year later by Antoniadou and Chintiroglou (2005), on hard substratum sciaphilic assemblages (15-40 m) in the North Aegean.
In the Greek seas, Brachiopoda have been recorded in habitats ranging from dimly lit hard substrata of the sublittoral zone (e.g. caves and coralligenous assemblages) to bathyal muddy bottoms and deep-water coral facies.
Overall, the brachiopod fauna of the Greek seas comprises 85% of the Mediterranean brachiopod species, being considerably richer than in the other countries of the eastern basin: 5 species have been reported to date from Turkey (Çinar 2014, Gönülal andGüreşen 2014), Cyprus, and Lebanon, and 2 species from Israel (Logan 1979, Logan et al. 2002. No endemic bachiopods have been found in the Greek seas. Nevertheless, given the low number of studies focusing on dark habitats in the area, further research is expected to increase our knowledge on the brachiopod fauna of the eastern Mediterranean.