Feeding ecology in sea spiders (Arthropoda: Pycnogonida): what do we know?

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Feeding ecology in sea spiders (Arthropoda: Pycnogonida): what do we know? Lars Dietz1,2*, Jana S. Dömel3, Florian Leese3, Tobias Lehmann4,5 and Roland R. Melzer4,5,6

Abstract: Sea spiders (Pycnogonida) are a widespread and phylogenetically important group of marine arthropods. However, their biology remains understudied, and detailed information about their feeding ecology is difficult to find. Observations on pycnogonid feeding are scattered in the literature, often in older sources written in various languages, and have never been comprehensively summarized. Here we provide an overview of all information on feeding in pycnogonids that we have been able to find and review what is known on feeding specializations and preferences in the various pycnogonid taxa. We deduce general findings where possible and outline future steps necessary to gain a better understanding of the feeding ecology of one of the world’s most bizarre animal taxa. Keywords: Pantopoda, Marine arthropods, Food chain, Benthos, Community ecology

Background Sea spiders (Pycnogonida) are a phylogenetically distinct group of marine arthropods with about 1500 species. General reviews of their biology were provided by King [1] and Arnaud & Bamber [2]. Almost all species have a holobenthic lifestyle. They are particularly abundant and species-rich in the polar regions, where genetic studies have identified several cases of unrecognized diversity [3, 4]. Although pycnogonids are widespread in all oceans and have been known to science for over 250 years, the feeding habits of most taxa remain poorly studied and a detailed review on the feeding ecology of pycnogonids has, to our knowledge, never been published. Observations on this topic are generally scattered throughout the literature, and especially publications written in languages other than English are often difficult to find. General textbooks usually only state that pycnogonids feed mostly on sessile prey, such as coelenterates, sponges and bryozoans (e.g., [5]). In the present paper, we review all available observations published in the last two centuries including both * Correspondence: [email protected] 1 Zoological Research Museum Alexander Koenig, Statistical Phylogenetics and Phylogenomics, Adenauerallee 160, D-53113 Bonn, Germany 2 Faculty of Mathematics and Natural Sciences, University of Bonn, D-53012 Bonn, Germany Full list of author information is available at the end of the article

detailed studies and preliminary notes, thus providing a state of the art summary of known food preferences for this bizarre and highly understudied group of exclusively marine arthropods. Additionally, we discuss morphological correlates of different feeding preferences and the occurrence of generalism vs. specialization in various pycnogonid taxa.

Morphological features for food uptake A pycnogonid that features all appendages used for feeding (Nymphon gracile) is pictured in Fig. 1c. As the main organ for food uptake, pycnogonids have a unique triradially symmetric proboscis with