The Malacologist 

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## NUMBER 77                                                            AUGUST  2021 

## **Contents** 

Editorial ………………………………………………… Notices …………………………………………………… Invited article ………………………………..……..….. Council of the Malacological Society of London taking part in the virtual AGM ………...……. Early Career Research Grant Reports 

AGM Composition of Council ………………………………... AGM Conference— _Molluscs in Extreme Environments_ 

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….. …………………………………... Book and film reviews ………………..…….... ………………..……....………………..…….. Annual general meeting—spring 2021 Annual Report of Council 


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•  ....................<br>in memoriam<br>..............................................<br>..............................................<br>...........................................<br>Forthcoming Meetings<br>..........................................................<br>Grants and Awards Of The Society.............................<br>Membership Notices …………………………………….…..<br>**----- End of picture text -----**<br>




**Despite covid-19, malacologists make their best endeavours. The mages above are from the Early Career Research Grant report of Alice Wilson McNeal on mpacts of ocean acidification and pharmaceutical contamination on** _**Mytilus edulis**_ **(see report on page 21).** 

The Malacological Society of London was founded in 1893 and registered as a charity in 1978 



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**EDITORIAL** 

It is still the time of covid. As I write, English case numbers stand at a mean of 31698/day  and deaths at a mean of 100/day over the last week.  That is _circa_ 50000 extra deaths per year (compare this with approximately 3000 mortalities when the Twin Towers  were felled in New York). While the hospital situation seems to have improved since the last issue of _The Malacologist_ , many young and some old people are developing long covid, while the country struggles to keep on a ‘normal’, even keel, including for those pursuing the objectives of malacology, such as running meetings. Despite the difficulties, the President John Grahame hosted an excellent virtual meeting of the **AGM** and a **virtual conference on** _**Molluscs in Extreme Environments,**_ held on the same day. The abstracts from the conference are presented on pages 34-43 this issue. This issue also presents an invited article on the biology of the molluscan shell. This was my personal indulgence, since I had encountered the author, Alessia Carini, when she was a presenter in a quickfire session at the 2020 Forum.  Some fifty years ago, I had researched the biochemistry of  freshwater molluscan shells, so I was particularly intrigued to see how things had developed in the interim. 

It is encouraging to the Society’s efforts to be able to present several **Early Career Research Grant Reports,** including an account of Lukas Ostermair’s work on _Molecular analyses of solenogaster midgut contents to determine food sources ,_ the report of  Alice Wilson-McNeal on the _Impacts of ocean acidification and pharmaceutical contamination on Mytilus edulis,_ and the report of Samuel Abalde on _Using shotgun sequencing for disentangling a taxonomic jumble: the case study of the skeneimorphs._ 

There are two **book reviews** —firstly of the book _Molluscan genomics: broad insights and future directions for a neglected phylum by Angus Davison and Maurine Neiman_ which summarises a recent Royal Society conference. Secondly there is a review of Peter Godfrey-Smith’s book entitled _Other minds—the octopus and evolution of intelligent life._ The review also references a recent BBC documentary on octopuses. 

Unfortunately, _The Malacologist_ also carries **obituaries.** Here there are tributes to Brian Morton, Arie W. Janssen and Jack Burch. 

Finally, this issue covers the formalities of the AGM, together with its associated virtual conference _Molluscs in Extreme Environments._ So, this turns out to be rather a large issue. Here’s hoping that all our readers stay covid free and above all, safe. 

Emeritus Prof. Georges Dussart Canterbury Christ Church University **TAXONOMIC/NOMENCLATURAL DISCLAIMER** Canterbury, This publication is not deemed to be valid for taxonomic/nomenclatural purposes Kent CT1 3JZ [see Article 8b in the International Code of Zoological Nomenclature 3[rd ] Edition georges.dussart@canterbury.ac.uk (1985), edited by W.D. Ride _et al_ .]. 

## **NOTICES** 

## **J Frances Allen Institute of Malacology Student Research Award** 

The Board of Directors of the Institute of Malacology, publisher of the academic journal _Malacologia_ , is pleased to announce the recipients of the 2021    J Frances Allen Institute of Malacology Student Research Award, a newly created award to support field or laboratory research on molluscs conducted by a student enrolled in a degree-granting program. We received 17 outstanding proposals from undergraduate and graduate students representing nine countries. The proposals were reviewed by a committee of the IM Board and were evaluated on the quality, feasibility, and potential significance of the proposed research. 

The first recipients of this award are: 

- Nicholas Gladstone, Auburn University, USA: Re-evaluation of the phenotypic plasticity hypothesis in a freshwater gastropod complex using phylogenomics, population genomics, and morphometrics (Pleuroceridae: _Lithasia_ ) 

- Noelia Sa nchez, National University of La Plata, Argentina: Conoidean gastropods from unexplored Argentine deepwaters: the family Drilliidae 

- Taro Yoshimura, The University of Tokyo, Japan: Sulfur detoxification by biomineralization in the hadal chemosynthetic bivalve _Axinulus hadalis_ (Thyasiridae) 

Please join me in congratulating the successful applicants. We wish them every success in completing their proposed research! 

Ellen Strong President Institute of Malacology 




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## **The Octopus in My House** 

This is the title of a BBC programme about the relationship between a Professor and diver  David Scheel and a wild octopus. 

ON BBC iplayer—BBC Two - Natural World, 2019-2020, The Octopus in My House 

See page 26 for a review of this programme. 


## **Squid in Space** 

Images, selected by the Bailey-Matthews National Shell Museum's Curator and Science Director Dr. Jose H. Leal convey a range of the singular beauty in the world of molluscs. For more information visit the website at www.shellmuseum.org/in-focus 

- The tiny squid illustrated below were due to head into space on June 3, 2021, along with many other scientific experi ments aboard SpaceX’s 22nd cargo resupply mission to the International Space Station. The squid were part of the UMAMI study which examines the effects of spaceflight on interactions between beneficial microbes and their animal hosts. UMAMI stands for Understanding of Microgravity on Animal-Microbe Interactions. Microbes play a significant role in the normal development of animal tissues and in maintaining human health. The Understanding of Microgravity on Animal-Microbe Interactions (UMAMI) study uses bobtail squid and bacteria to examine the effects of spaceflight on interactions between beneficial microbes and their animal hosts. This type of relationship is known as symbiosis. Beneficial microbes play a significant role in the normal development of animal tissues and in maintaining human health, but gravity’s role in shaping these interactions is not well understood. This experiment could support the development of measures to preserve astronaut health and identify ways to protect and enhance these relationships for applications on Earth. This investigation helps _Image Credit: Courtesy of Jamie S. Foster_ determine whether spaceflight alters the mutually beneficial relationship, which could support development of protective measures and mitigation to preserve astronaut health on long-duration space missions. The work also 

could also lead to a better understanding of the complex interactions between animals and beneficial microbes, including new and novel pathways that microbes might use to communicate with animal tissues. Such knowledge could help identify ways to protect and enhance these relationships for better human health and well-being on Earth as well. Yvette Smith May 26, 2021 

## **Using slug slime to engineer adhesives for diverse wet surfaces** 

_Abstract of the paper   -_ Adhesion to wet and dynamic surfaces, including biological tissues, is important in many fields but has proved to be extremely challenging. Existing adhesives are cytotoxic, adhere weakly to tissues, or cannot be used in wet environments. We report a bioinspired design for adhesives consisting of two layers: an adhesive surface and a dissipative matrix. The former adheres to the substratum by electrostatic interactions, covalent bonds, and physical interpenetration. The latter amplifies energy dissipation through hysteresis. The two layers synergistically lead to higher adhesion energies on wet surfaces as compared with those of existing adhesives. Adhesion occurs within minutes, independent of bloodexposure and compatible with _in vivo_ dynamic movements. This family of adhesives may be useful in many areas of application, including tissue adhesives, wound dressings, and tissue repair….. 

_From within the text_ - The design is inspired by a defensive mucus secreted by slugs ( _Arion subfuscus_ ) that strongly adheres to wet surfaces. This slug adhesive consists of a tough matrix with interpenetrating positively charged proteins. Our tough adhesives are made up two layers: (i) an adhe- 

sive surface containing an interpenetrating positively charged polymer and (ii) a dissipative matrix (Figure). The adhesive surface can bond to the substratum through electrostatic interactions, covalent bonds, and physical interpenetration, whereas the matrix dissipates energy through hysteresis under deformation. _Reference_ 

Li A., Celiz A.D., Yang J., Yang Q., Wamala I., Whyte W., Seo, B., Vasilyev N.V., Vlassak J.J., Suo Z, & Mooney D.J. 2017. Tough adhesives for diverse wet surfaces. _Science,_ **357** : 378–381. 




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## **OctoCam is LIVE!** 

On April 30th, the Bailey-Matthews Shell Museum unveiled its live stream of OctoCam - an underwater camera that gives visitors a unique view into the nocturnal life of its Giant Pacific Octopus. To see the octopus live, plus highlight footage of its very active moments follow:- 

www.shellmuseum.org/octocam Dr. Jose  H. Leal, Science Director and Curator Bailey-Matthews Shell Museum 3075 Sanibel Captiva Rd. Sanibel, FL 33957, USA 


## **Companions for the evening** 

Patricia Highsmith once went to a London cocktail party with a "gigantic handbag" that "contained a head of lettuce and a hundred snails" which she said were her companions for the evening. 


## **New book on South American bivalve molluscs** https://bit.ly/2HIPPj2 


Gene Coan and **Paul Valentich-Scott** have completed the eastern Pacific marine bivalve series (https:// bit.ly/2HIPPj2) and are making the combined bibliography for the three books available online. This includes over 7,600 references cited in their mongraphs. In addition, John Taylor and Emily Glover have shared the bibliography from their recent book on the Bivalve Family Lucinidae (https://bit.ly/3b1iSKg). This has added an additional 500 references to the bibliographies, including those on vent and seagrass communities. The database is accessible through an opensource freeware called Zotero (https://www.zotero.org). A free Zotero account should be set up if you wish to access this database. If you are interested in accessing this database, send your email address pvscott@sbnature2.org. You will need to send  the email you used to set up a zotero account. 

Paul Valentich-Scott, Curator Emeritus of Malacology Santa Barbara Museum of Natural History 2559 Puesta del Sol, Santa Barbara, CA 93105 www.sbnature.org 


## **Late evening work** 

The slugs and snails in my garden (particularly _Arion ater_ and _Cornu aspersa_ a) re bullies, homing in on weaker plants and stripping an individual plant of its leaves in a single night.  Starting in March, and for several weeks, I go into the garden before bedtime with a torch, a jam jar and a plumber’s glove, collecting slugs and snails from my seedlings. I put them in the freezer overnight to painlessly kill them, then bury the corpses deep in the compost heap. The first several  nights, I get a jam jar full each night.  The numbers decline as I make inroads into the population but I never get them all. Sometimes it seems there is the molluscan equivalent of a small sheep feeding at night on my flowers and veg. 

GBJD 




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## **The Malacological Society of London’s** _**Molluscan Forum**_ 

Because of Covid-19, the Molluscan Forum will take place in a different format this year. We still plan to hold the event and will do so via an online platform (probably Zoom) on Thursday 18t[h] November 2021. 

As in previous years, for those wishing to offer a presentation, please fill out the attached registration form. For those just wanting to attend, please let me know so I can keep track of attendees. The **deadline for registrations and talk applications is October 1st 2021.** Presenters will be informed of successful applications soon after **.** 


Registration is **free!** 

In lieu of posters, we will instead have 5 minute (3 slide maximum) **Quickfire PowerPoint** presentations, 

We hope these changes will still allow all those who wish to attend and present the chance to do so, even if we can’t offer the customary wine reception! 

For further details  and an application form, see page 51. 

The contact is Dr Phil Hollyman British Antarctic Survey, Cambridge CB3 0ET Email: phyman@bas.ac.uk 


## **A climate-warming relevant snail joke from Tim  Pearce** 

https://www.facebook.com/watch/?v=796957347660223 


## **Molluscan genomics: broad insights and future directions for a neglected phylum** 

Royal Society Publishing has recently published a special issue of Philosophical Transactions B - _Theo Murphy meeting Molluscan genomics: broad insights and future directions for a neglected phylum_ organized and edited by Dr Angus Davison  and Dr Maurine Neiman. Articles can be accessed directly at www.bit.ly/PTB1825. If your institution does not give access to the full text, corresponding authors will probably be happy to send a pdf, or else contact Maurine or Angus. A print version is also available at the special price of £35.00 per issue, by contacting Debbie.vaughan@royalsociety.org Dr Angus Davison, Reader and Associate Professor in Evolutionary Genetics School of Life Sciences, University Park University of Nottingham Nottingham, NG7 2RD angus.davison@nottingham.ac.uk 

(This Royal Society special issue is reviewed on Page 25 of  this issue of _The Malacologist_ ) 


**Spot the mollusc** 





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## **Bailey-Matthews National Shell Museum** 


## **The Chiton and the Whelk** 

Museum's Senior Aquarist Carly Hulse took this great photo, at the Cold-water Touch Pool, of  a Lined Chitons ( _Tonicella lineata_ ) riding a Kellet’s Whelk ( _Kelletia kelletii_ ). The chiton needs hard substrata to live and feed on, and the whelk’s shell serves that purpose. 


**Lined Chiton riding a Kellet's Whelk. Photo by Carly Hulse.** 

The whelk’s right tentacle is touching the bottom of the aquarium. All molluscs in the Cold-water Touch Pool  were obtained from the West Coast of the US, where the water temperature is in average at least several degrees cooler than the water temperature in Southwest Florida. 

(with the permission of Jose  H. Leal, Bailey Matthews National Shell Museum) 


## **Mollusc conservation—Tentacle** 

The latest issue - number 29 - of Tentacle (the newsletter of the IUCN - Species Survival Commission - Mollusc Specialist Group) is now available on the web. 

http://www.hawaii.edu/cowielab/issues.html 

Tentacle is an on-line only publication, though of course you may print it for yourself if you wish. 



**Malacological cosmetic treatment as portrayed in the new TV series ‘Murder in Mallorca’** 


**Mollusc themed dinner plate from the Charente region of France** 




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## **Article on the co-existence of** _**Crassostrea**_ **and** _**Mytilus**_ 

Ecological impacts caused by invasive alien species can be severe but may vary depending upon environmental conditions. Many European populations of the native mussel, _Mytilus edulis,_ have been invaded by the Pacific oyster, _Crassostrea (Magallana) gigas_ . Although widespread invasions have occurred, interactions between _M. edulis_ and _C. gigas_ have largely been investigated with regards to competition for space and food as well as effects on species assemblages. Experimental investigation of competitive interactions on physiological responses of the two species requires further exploration. To this end, we used a 12-month field manipulation experiment to examine growth rates, mortality and condition indices of the two species occurring in monospecific and heterospecific groups. Growth rates and mortality of both species were similar in monospecific and heterospecific groups, whereas condition indices were significantly reduced for both species in heterospecific groups. Growth rates and condition indices also differed amongst experimental sites, potentially due to differing water motion. Shell weight-length relationships did not explain the observed differences in condition for either species. Coexistence between the two species may occur but could be detrimental for both species. A preliminary viewpoint that water motion can mediate competitive interactions between these species is also provided. 

P. Joyce, D. Smyth, J. Dick  & L. Kregting  (2020) 

Coexistence of the native mussel, _Mytilus edulis_ , and the invasive Pacific oyster, _Crassostrea (Magallana) gigas_ , does not affect their growth or mortality, but reduces condition of both species 

_Hydrobiologia_ https://doi.org/10.1007/s10750-021-04558-1(0123456789().,-volV)( 01234567 


## **Mollusc of the Year** 

This award was organised by the Senckenberg Research Institute, together with the  LOEWE Translational Biodiversity Genomics centre, and Unitas Malacologica. There were over 120 nominations for any molluscan species from any marine, terrestrial, freshwater zone or from any country. The first, ever, international Mollusc of the Year award went to _Argonauta argo._ More than 21,000 votes were cast by the public, with _Argonauta_ capturing 36% of the vote, followed closely by _Polymita pita,_ and _Congeria kusceri_ (see image below). Specimens of _Argonauta argo_ from the nominating group, at Stazione Zoologica Anton Dohrn, Naples, Italy, have now arrived at the Senckenberg LOEWE Translational Biodiversity Genomics centre, where a team  will sequence the complete genome during 2021. The organisers send a huge’ thankyou’ to all who nominated species, and all who voted! It is hoped to repeat the award next year. There is more about the competition here: https://tbg.senckenberg.de/mollusc/ Julia Sigwart 



## **Decapitated sea slug grows a new body** 

Video at:-instagram.com/p/CMNQ_t0qOx7/?igshid=mn8no78cwweoscitechdaily.com Sayaka Mitoh & Yoichi Yusa, 2021,  Extreme autotomy and regeneration of the whole body in photosynthetic sea slugs. _Current Biology_ , **31** : 5R233–R234, **DOI: 10.1016/j.cub.2021.01.014** 




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## **Cuttlefish exert self-control** 

The ability to exert self-control varies within and across taxa. Some species can exert self-control for several seconds whereas others, such as large-brained vertebrates, can tolerate delays of up to several minutes. Advanced self-control has been linked to better performance in cognitive tasks and has been hypothesized to evolve in response to specific socioecological pressures. These pressures are difficult to uncouple because previously studied species face similar socioecological challenges. Here, self-control and learning performance is investigated in the cuttlefish, an invertebrate that is thought to have evolved under partially different pressures to previously studied vertebrates. To test self-control, cuttlefish were presented with a delay maintenance task, which measures an individual’s ability to forgo immediate gratification and sustain a delay for a better but delayed reward. Cuttlefish maintained delay durations for up to 50–130 s. To test learning performance, a reversal-learning task was used, whereby cuttlefish were required to learn to associate the reward with one of two stimuli and then subsequently learn to associate the reward with the alternative stimulus. Cuttlefish that delayed gratification for longer had better learning performance. These results demonstrate that cuttlefish can tolerate delays to obtain food of higher quality comparable to that of some large-brained vertebrates. 

Schnell A.K., Boeckle M., Rivera, M., Clayton N.S. & Hanlon R.T. 2021.     Cuttlefish exert self-control in a delay of gratification task. _Proc. R. Soc. B_ **288** : 20203161. https://doi.org/10.1098/rspb.2020.3161 



## **Slug (** _**Lehmannia nyctellia**_ **) slides down a mucus thread like a spider on silk** 

Published:03 March 2021https://doi.org/10.1098/rspb.2020.3161 

https://www.iflscience.com/plants-and-animal /World-First Sighting Of Slug Sliding Down Mucus Thread Like A Spider On Silk | IFLScience 




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## **Call for Applications: 2022 Conservation Leadership Programme (CLP) Team Awards** 


## **Funding opportunity for young scientists working in species conservation** 

The funding comes with a training opportunity for one team member to take part in a highly regarded course, usually held in Cambridge UK, although recently it has been held online. 

Previous grants have tended to be awarded to bird, mammal and plant projects, so the competition is high. However, with a good project and clear outcomes, there is an chance for young invertebrate conservationists to make an application for a place. Projects working to resolve understanding of the status of _Data deficient species_ can be equally valid as conservation assessments and recovery plans for threatened species. Freshwater spring snails, Cave invertebrates or species that exist in world heritage sites might be good candidates for projects. 

- - - https://www.conservationleadershipprogramme.org/our projects/latest projects 2021/ 

Successful candidates have the opportunity for follow-up funding for projects. 

The Conservation Leadership Programme (CLP) is now accepting applications for our 2022 Team Awards. Future Conservationist Awards are worth up to $15,000 each and support projects in low- and middle-income countries and some highincome islands in the Caribbean and Pacific. Funding enables early-career conservationists to conduct scientific research, encourage and promote pro-conservation attitudes to better conserve and manage the natural world. Each project must have at least three members and all team members must have less than five years of paid conservation experience. Projects can last 3-12 months and must focus on a threatened species. In addition to project funding, CLP will invite one member of each award-winning team to their international training course where they will gain a range of conservation management and leadership skills as well as build their professional network. 

## **The application deadline is 10 October, 2021.** 

For more information, including the eligibility criteria and detailed application guidelines, please visit our website. Applications must be submitted via the online platform (potential applicants will need to register to be able to complete the application form). Contact clp@birdlife.org with any questions. 

Stuart.Paterson@fauna-flora.org> 


## **16th Deep Sea Biology Symposium, Brest, France     12-17 September 2021** 


The meeting includes special sessions on the impacts of deep sea mining and polymetallic mining. Programme  https://wwz.ifremer.fr/16dsbs/Program Conference information       https://wwz.ifremer.fr/16dsbs/Conference-Information 




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## **New book—Bivalve seashells of Western South America** 



## **Carnivore Dilemma** 


Most bivalve molluscs are filter-feeders, straining water to get their food, usually microscopic plants. Exceptions to this include marine clams that prey on small creatures, such as crustaceans. Some of these clams, the septibranchs, use a flexible, muscular "wall", or septum, as a diaphragm pump that allows the animal to quickly and decisively suck prey in. The inch-long Specter Clam, _Dilemma spectralis_ Leal, 2008 (below), a deep-sea dweller found off Vanuatu in the SW Pacific, is one of them. It is possible, given that _Dilemma_ clams live permanently attached to rocks, that Specter Clams may use some form of chemical attraction to lure nearby prey. 

Fig. 2 shows from left, a whole preserved clam, the clam with shell removed, and drawing (by Kimberly Nealon) showing the stomach with its crustacean food, in this particular case an isopod crustacean. The drawing also shows the _incurrent siphon_ , or opening through which prey is ingested. 

**Fig. 1** _**Dilemma spectralis**_ **, different views of an unusual shell. From the original species description.** 


Jose H Leal (2008) A remarkable new genus of carnivorous, sessile bivalves (Mollusca: Anomalodesmata: Poromyidae) with descriptions of two new species. _Zootaxa_ **1764** :1-18 

**Fig. 2** 




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## **Marine shells: the beauty and the resilience** 

**The Swire Institute of Marine Science and School of Biological Sciences** , The University of Hong Kong, Pokfulam Road, Hong Kong SAR Email: acarini@hku.uk 

People strolling along the beach have been collecting shells since prehistoric times and treasured them as beautiful, sometimes spiritual, marine tokens (Scales, 2015). Many public beachgoers who stumble upon empty shells do not realize that these are the washed-up legacy of living organisms. This could be because marine shells can resemble perfect pieces of elegant, stony, small (and occasionally large) rocks. They are indeed composed of a mineral analogous to limestone, yet, contrastingly, they can be shaped in extraordinary forms, and, as you know if you have ever  cut your foot on the beach, of puncturing sharpness and strength. Marine shells recovered on beaches are often either the coiled exoskeletons of gastropods or the disjointed half of bivalve shells, and here I will mostly draw examples from the latter group. 

Molluscs are the second species richest phylum in the world after the Arthropods and the class _Bivalvia_ accounts for an estimated 10,000-20,000 extant species (Gonzalez _et al_ ., 2015). They are characterized by two tightly fastened valves, sometimes but not exclusively, symmetrical. Since their appearance in the Cambrian, around half a billion years ago, they have spread to an incredible number of different habitats, from the tropics to the arctic, from lakes to hydrothermal vents, and survived several catastrophic events in our Earth’s history such as the Palaeocene-Eocene Thermal Maximum (PETM). The PETM was characterized by warming, acidification and hypoxia that lead to severe extinctions and changes in global ecosystems. Little lasting impact was sustained by molluscs however (Ivany _et al_ ., 2018). Simply, these unpretentious invertebrates are resourceful. The evolutionary success of the bivalves is the manifestation of small modifications to an already simple animal body plan (Plazzi _et al.,_ 2011) and, importantly, the reason bivalves are traditionally thought to have been so successful is the secretion of the external shell (Clark _et al_ ., 2020). 

The bivalve shell is a composite biomineral, incorporating mineral and organic components, and molluscs produce the highest diversity of biominerals of any animal group (Lowenstam & Weiner, 1989). The mineral of choice by the vast majority of bivalves is calcium carbonate (CaCO3), which, in its abiotic form is typically unimpressive, not only in shape but also in mechanical properties. However, when moulded into a shell by a mollusc, it takes beautiful, complex, and mechanically superior forms. This is a good example of structural potential once biology and evolution join forces to create forms able to persevere for millions of years. 

Evidence suggests that the molluscan shell evolved from an ancestor with a chitinous scaffold (Murdoc, 2020). Since eukaryotic cells have the inherent capacity to bind calcium ions and control their internal carbonate chemistry, accumulating ions for calcification simply exploited some ancient cellular features (Knoll, 2003). However, biomineralization is energetically costly, hence, there are metabolic constraints on organisms that limit the evolutionary potentialfor skeletal components (Knoll, 2003; Schoeppler _et al.,_ 2019). 


**Fig 1. Some of the bivalve shells in the  collection in the Hong Kong Biodiversity Museum. Shells are not to scale.** 

Even so, for biomineralization to be so widespread, the benefits of having a shell must offset the metabolic costs, yet the energy spent to secrete a shell must be flexible as a response to the requirements by the external environment (Knoll, 2003). Ultimately, the evolutionary and metabolic constraints are the drivers for the diversity of structures that we see today. Intuitively, the bivalve shell functions as armour and structural support for the vulnerable, and, regrettably for the bivalve, tasty soft body for a predator. Nonetheless, bivalve shells can have several different functions other than structural ones. 

CONTINUED> 



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## CONTINUED> 

For example, several bivalve species live in intertidal regions where they are exposed to air or sediments for extended periods. In these scenarios, the shell prevents dehydration by locking tightly and retaining water. In some cases, shells can act as reservoirs and release ions to neutralize internal pH which quickly decreases due to excessive production of metabolic 

CO2 (Lowenstam 1981). 


**Fig. 2. Extract from Fremy, E. 1855. Roughly translated, Fremy reports that certain shells, when treated with acid, release a remarkable organic matter, of a shiny and felted appearance, insoluble in water, alcohol, and ether, capable of withstanding the extended action of acids. He names it conchiolin and then describes its organic composition This was probably the first ever description of conchiolin.** 

The basics of shell formations are that calcium, carbonate and/or bicarbonate ions are extracted from seawater and concentrated in a confined space by the bivalve at a much higher level than the surrounding seawater. Isolation of this medium from the external seawater is necessary to facilitate and promote calcification. CaCO3 is then deposited together with an organic fraction called the shell organic matrix. This matrix is extracellular, and it is typically composed of organic compounds such as polysaccharides (mostly chitin), proteins, and lipids. Pioneering  research on bivalve shells and their organic matrices has been done since at least the 1800s. Other than geologists and palaeontologists, few chemists found interest in this biomineral . An exception was  Edmond Fremy (1855), who, in his quest to describe as many as possible biominerals from the animal kingdom,  came across molluscan shells, thanks to a collaboration with a certain M. Valenciennes (Fig. 2). After applying his standard decalcification protocol to marine shells, 

he obtained an insoluble, shiny organic substance he named _conchiolin_ after its similarities with ossein. Conchiolin had, however, a different chemical behaviour from ossein, and he also points out the presence of a carbohydrate, chitin, a trait he found shells’ share with arthropod exoskeletons and cnidarian skeletons. This is also one of the first times that researchers found how much these invertebrate exoskeletons have in common with human skeletons, a parallel that is still an active field of research (Loh _et al.,_ 2021, Clark _et al.,_ 2020). 

A lot of shell literature followed the work of Fremy, although some of this material is now difficult to access and often requires translations from different languages. Fast-forwarding to the 20[th] century, we see that renewed, international interest was developed in the 1960-80s. While the presence of an organic matrix within shells was well established, we see more and more attempts to characterize it in increasing detail. It was known that a large part of the shell organic matrix was composed of proteins, and with their amino acid characterization, Weiner & Hood (1975) provided evidence that shell proteins are rich in negatively charged amino acids. Weiner & Traub (1980) with their improved x-ray analysis of the insoluble protein fraction of the matrix confirmed that it displays similarities with silk fibroins and has a β-sheet conformation, probably in an antiparallel fashion. As new instruments emerged and were more widely accessible, Weiner, Talmon & Traub (1983) published the first analysis of organic matrices with electron diffraction patterns. Strikingly, the observations from these and many concomitant experiments, still make up a lot of the ground theory and assertions in the biomineralization field. At the same time, biomineralization scientists started to come together since at least 1970, when the first international symposium was held in Germany. An account of the progress since their second international meeting can be found in the Science bulletin from 1977, where Towe paints a rather negative picture (Fig. 3). 

No overlaying mechanism is observed, and scientists are baffled by the immense number of structures, their complexity, and the absence of an observable common underlying mechanism. Recent biomineralization review papers similarly state that, despite the newest developments in the field, the complex variety of three-dimensional shell morphologies is largely unexplained (Clark _et al_ ., 2020). While this is true, the fact is that the field of biomineralization has greatly grown, several scientific groups from all over the world revolve around it, innumerable datasets have been collected and ever more structures have been investigated. Flourishing information and the development of ever more advanced equipment has led to even further investigations opening new debates and reshaping questions. 

The main organ in charge of the complex biomineralization apparatus is the mantle. The mantle is a relatively thin tissue that overlays both valves and stretches as needed to tackle multiple tasks within and outside of shell formation. There is also evidence that other cells could be involved in biomineralization in different species (see for example Sillanpa a _et al_ ., 2016), however, the mantle remains the best  studied. Gene expression analyses of mantle tissues have greatly increased the amount of information we possess on molluscan biomineralization. Nevertheless, the picture remains puzzling, and this is in part due to the enduring neglecting of molluscan genomes as well as the fast evolutionary speed of biomineralization genes which make phylogenetic comparisons difficult especially when no dominant molluscan model species has yet emerged. However, a renewed effort is now being put into expanding our databases and more information is flowing in, thanks to international collaborations (Davison & Neiman, 2021). It has been repeatedly observed that there are high rates 

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**Fig 3. Review of papers published after the second biomineralization symposium (Georgetown, USA, 1974). Printed in** _**Science**_ **196 (4296) pp 1311.** 

of lineage-specific gene co-option and loss from mantle tissues, rapid evolution of coding sequences (Aguilera, McDougall & Degnan, 2017), and extensive duplication of shell matrix protein families (Miyamoto _et al.,_ 2013, Takeuchi _et al.,_ 2016). Since molluscs produce the highest diversity of biominerals of any animal group, the evolvability of the mantle could provide a molecular explanation for this (McDougall & Degnan, 2018). Whether this confers any selective advantage to molluscs, however, or whether it is a byproduct of the molecules necessary for shell assembly, is not known (Kocot _et al.,_ 2016). Exploring these questions would be key to understanding the adaptation potential for this group in the face of rapid environmental change. 

The complexity behind shell formation is further enriched when we include increasing evidence for epigenetic post-transcriptional regulation of biomineralization processes through microRNAs (Zheng _et al.,_ 2016) and long noncoding RNAs (Zheng _et al_ ., 2020). In addition, the Mediterranean mussel genome has been reanalysed and has been found to contain an unexpectedly high number of dispensable genes for an animal (Gerdol _et al.,_ 2020). Dispensable genes are characteristic of microorganisms such as bacteria, and they are correlated with the ability to respond to selection and inhabit new niches, something that bivalves, which (1) survive in extreme environments, (2) are often exposed to highly fluctuating environments and (3) can be cosmopolitan, have in common with microorganisms.  This feature is likely to be found in other bivalves with invasive species traits. All of these observations are important to link the thought of a shell as a solid, rock-like immutable structure and the fact that a great diversity of bivalve and molluscan shells exists, not only between but also within species. For example, the plasticity of shell formation is particularly evident in certain groups such as cementing oysters that create shells so polymorphic that often, species identification based on shell morphology is impossible (Xia, Yu & Kong, 2008). Contrary to all predictions, mussels have been reported to quickly modify their shell structure to adapt to changes in the environment and predation pressure (Telesca _et al_ ., 2020). It has also been suggested that rapid responses to environmental stress are facilitated by maintaining high genetic diversity in the brooding population (Bitter _et al.,_ e2019). In fact, mussel larval shells size can be sustained even in very low pH seawater through fitter genotype and phenotype selection (Bitter e _t al.,_ 2019). Therefore, the lack of observations of a clearly defined biomineralization genetic tool and common shell architecture, even within closely related species, is not all that surprising. Clearly, their genetics offer a unique of opportunities for us to learn about plasticity and evolutionary biology. 


**Fig. 4. Adult hard clam shell (** _**Meretrix**_ **sp.). A) Bare valve after removal of the periostracum, B) valve with original periostracal cover and C) shell ligament.** 

The making of the bivalve shell begins very soon after fertilization. Bivalves are spiralian animals that undergo unequal embryonic cleavage until bilateral cleavage of dorsal cells triggers the establishment of the unique doubling of the dorsal shell (Kin _et al.,_ 2009). It is believed that the bivalve cleavage pattern is highly conserved, and that canalisation of development is key to the establishment, and maintenance, of the bivalve body plan (Kin _et al.,_ 2009). A series of cell signalling will initiate the formation of the so-called _shell field—_ an early organic matrix, 

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**Fig. 5. Example of a bivalve larval shell imaged using scanning electron microscopy. A) Entire Hong Kong oyster larval shell. B) Details of the larval ligament structure from outside the shell and C) view from inside.** 

which flattens the trochophore larva into a version of the bivalve body frame with which we are familiar (Fig. 5A).  Mineral deposition co-occurs upon the organic matrix and at high rates, relatively much higher than in adult counterparts. The bivalve larva mass has been observed to be 90% shell which needs to be deposited by relying mostly, if not exclusively, on the energy provided by the maternal oocyte (Waldbusser _et al.,_ 2013). The first bivalve larval shell, which is called a D-shell, is similar among different species with very divergent adult shell morphology. Differently from the settled, mature versions of the bivalve shells with which we are familiar, the larva in its shell is microscopic, free-swimming, and in many species, fully planktonic. Despite all these differences and metabolic challenges, the larval shell already includes all of the features that will also make up the adult shell: distinct layers of CaCO3, hinge ligament, periostracum, and a diverse organic matrix proteome. 

The periostracum and the ligament are hardened organic layers, mostly proteinaceous. These structures are visible in the first larval shell (Fig. 5 and 6) and are retained and further developed in the adult counterpart (Fig. 4). Ligament formation is initiated by specialised cells during larval development after valve partitioning (Moue za _et al.,_ 2006) and supports bivalve motility throughout its life. The ligament maintains its elasticity while keeping the valves tightly sealed yet flexible, and is a composite of crystals and an unusual combination of proteins (Suzuki _et al.,_ 2019). Similar to a tough insect cuticle, these structures are sclerotized, probably through quinone tanning of soluble proteins (Waite, 1977) and they are produced by the mantle like the rest of the bivalve shell. By contrast with the ligament, the periostracum is not mineralized, although intraperiostracal calcification is not uncommon in adult bivalves (Checa & Harper, 2014). As most clearly observed by transmission electron microscopy, the periostracum encloses the newly formed larval shell from hinge to mantle edge and it is the first observable part of the 

larval shell (Moue za _et al_ ., 2006). One of the prime functions of the periostracum is to provide waterproof support and a substratum for the growth of the outer shell layer (Taylor & Kennedy, 1969). An external organic layer is necessary to isolate the calcification space to create that highly concentrated fluid for CaCO3 deposition and to avoid immediate shell dissolution. The periostracum layer is the key player in more corrosive waters such as polar, subpolar, and low-salinity environments (Telesca _et al.,_ 2019). In these habitats, mussels within the same species modulate shell formation by secreting a thicker periostra- 


**Fig. 6. Bivalve larval shell example imaged using scanning electron microscopy. A) View inside a larval shell valve. B) Details of the larval periostracal cover and C) view of larval shell layers from a broken shell fragment.** 

cum and a less calcified shell as compared with their lower latitude relatives (Telesca _et al_ ., 2019). 

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Another adaptation is to increase the organic component within the shell itself. It has been suggested that shells have become more calcified and less organic over time, possibly due to the high metabolic cost of secreting the organic matrix (Palmer, 1992). Therefore, this is not necessarily the best approach to produce more functional shells, at least not on an evolutionary time-scale. It was also proposed that the introduction of calcite to replace aragonite was a cheap innovation for protection from dissolution as _calcite_ is less soluble than _aragonite._ These are the two main CaCO3 polymorphs assembled by bivalves: aragonite is harder and stronger, but it is also more soluble than the “cheaper” calcite. It is widely accepted that the primitive molluscan shell was wholly aragonitic (Taylor, 1973). While the bivalve larval shells are all still composed of aragonite, many adult bivalves, such as mussels, include a calcitic layer right under their periostracum. Since this layer would be more at risk of interacting with seawater especially during early deposition, it could indeed be beneficial against dissolution. Whilst aragonite is a more soluble polymorph, there is little evidence that aragonite layers are more soluble than calcitic ones in cold waters (Harper, 2000), but several studies on the effect of decreased pH show a more pronounced negative effect on aragonitic layers as compared to calcitic ones (reviewed in Byrne & Fitzer, 2019). It has been argued, therefore, that aragonitic shells were difficult to maintain and secrete during highly corrosive ancient calcite seas and that during these times the acquisition of a calcitic shell layer was advantageous (Harper, Palmer & Alphey, 1997). Reversing mineralogy is a costly business (Porter _et al_ ., 2010) and, therefore, the layer might have been maintained since. When rapid calcification is required, as during embryonic development or new edge growth, granular homogenous structures are observed (Schoeppler _et al_ ., 2019). As the granular layer grows, the number of crystal nucleation centres decreases and the CaCO3 layers transition in a predictable manner to columnar and then nacreous following a directional solidification model (Schoeppler _et al_ ., 2019). Nacre, mother of pearl, is easily the most commonly known structure, for its beautiful appearance and commercial value (Fig. 7). Not surprisingly, it has also been the model biomineralization structure in molluscan shell studies. A well-known nacreous material is also the oyster pearl, which is essentially an inside-out shell, with the periostracum at its centre (McDougall _et al_ . 2021). Pearls are arguably the most impressive piece of biomaterial produced by molluscs, and the most valuable (at least from a human perspective). Other than its beautiful appearance, nacre is known for being the strongest of all molluscan microstructures (Checa, 2018). Nacre’s architecture can absorb 1–3 times more mechanical energy than prismatic calcite and monolithic aragonite before total failure (Gim et al., 2019). Nacre also contains a large proportion of organics, compared to other layers, and these can help restrict crack propagation both within and between tablets, sustaining the overall architecture and allowing further structural loading (Gim _et al_ ., 2019). 

Similar sequences of different shell layers are found in calcitic oyster shells, from outer prismatic to the inner foliae layer which is an analogue to nacre. Nacre has evolved multiple times in different lineages (Jackson _et al_ ., 2010) following the _directional solidification_ model. These microstructural sequences, based on 

**Fig. 7     Mother-of-pearl comprising nacre** 

thermophysical, and chemical laws, appeared repeatedly during evolution (Checa, 2018: Schoeppler _et al.,_ 2019). An exemple is that calcitic foliae have developed independently in bivalves and gastropods, even though they form a weaker structure than nacre (Checa, 2018). While the large number of microstructures produced by molluscs is impressive, how have bivalves created just the right environment, for their assembly if the CaCO3 abiotic version is so uncomplicated and seawater conditions unfavourable? The answer seems to be that the physical and chemical boundaries for shell secretion are controlled by the direct action of the mantle and then remotely through the secretion of the extracellular conchiolin. 

As observed by Fremy in 1855, molluscan shell conchiolin includes a polysaccharide, chitin. Chitin is frequently used by organisms for structural support and probably contributes to the mechanical strength of the shell (Chan _et al_ . 2018). Not all shells show evidence of chitin however, and therefore, the abundance and role of chitin in molluscan shells have been debated. When present, it associates with proteins and the mineral phase. For example, in the pearl oyster, crystal defects become larger as chitin fibres are degraded by chitinolytic enzymes (Kintsu et al., 2017), and in the Eastern oyster when the mineral phase or proteinaceous fractions are dissolved, the chitin fibrils growth is stunted (Chan _et al.,_ 2018). There is evidence for chitin presence in the bivalve larval shell as well, where it strongly interacts with proteins (Weiss & Scho nitzer, 2006). 

Shell matrix proteins have received the most attention in recent years, probably due to advances in mass spectrometry. Models based on molluscan nacre proposed that the bulk of shell proteomes is composed of silk-like proteins with gel-like properties that are interspersed within the mineral tablets (Addadi _et al_ ., 2006). Indeed, several protein sequences similar to silk have been extracted from bivalve shells and often possess chitin-binding domains. It is reasonable to conclude therefore, that silk-like proteins and chitin provide structure to the matrix and the shell. In addition to silk-like proteins, classical models also include negatively charged proteins which have long been hypothesised to interact with positively charged calcium ions and therefore playing an active key role in biomineralization CONTINUED> (Weiner & Hood, 1975). 



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In addition to these components, it is well established that shell proteomes are characterized by repetitive, low complexity domains that have a biased composition and evolve rapidly by continual recruitment, deletion, and duplication (McDougall, Aguilera & Degnan, 2013). 

Most studies regarding the characterization of the shell organic matrix have focussed on nacre. We know from several proteomic studies however, that different mineral layers can have large differences in their shell proteomes (eg. Marie _et al.,_ 2012). Further, shell proteomes vary strongly with ontogeny (Zhao et al., 2018) and environmental fluctuations (Arivalagan _et al_ ., 2020). While proteins are estimated to only account for 1-5% of the shell weight, shell proteomes are quite diverse, and proteomics studies have accumulated a high number of sequences that often do not trace back to any known protein or function. However, functional studies have provided evidence for some long-hypothesized functions of soluble SMPs such as crystal nucleation (Feng _et al.,_ 2009), CaCO3 phase determination (Suzuki _et al.,_ 2009), initiation and inhibition of calcification (Sikes _et al.,_ 2000) and controlling crystallization patterns (Checa _et al.,_ 2016). Other than I improving the structural properties of bivalve shells, playing an active role in calcification and bivalve shell evolution, the organic matrix moulds the bivalve shell into a biochemical barrier for the animal. The identification of shell proteins with immunology-related domains used to be troubling, as they are associated with cellular functions and, were therefore, thought to be contaminants. The frequent identification of proteases, metalloproteinases, and protease inhibitors from shell proteomes, even after harsh biomineral cleaning, has now produced new scope for shell studies in the light of biochemical defence against pathogens. These expanding datasets and observations require new, updated, comprehensive models for the bivalve shell matrix that, while consolidating the traditional, well-established knowledge, also appreciates the novelty and nuances of newer findings. 

The fact remains that after decades of research, we are still baffled by these invertebrates’ resilient creations. The mechanical superiority of these shells offers great opportunities for us to create new bioinspired materials (Clark _et al.,_ 2020) and to explore their value for human health, with applications in wound healing, tissue engineering, and bone regeneration (Loh _et al.,_ 2021). Bivalves, and in particular their shells, are ecosystem engineers as they reshape their environment and create new homes to countless creatures while also providing an effective natural carbon sink (Fig. 8). 

Clearly, bivalve shells are more than just relics on the beach, but issues such as overconsumption, habitat loss, pollution, and climate change, to name a few, put at risk many of these species and their highly specialized shells. Even if it were not for our own consumption, health benefits, or ecological roles, understanding  shell formation is important to the safeguarding of the incredible shell diversity that has fascinated geologists, biologists, chemists, and the public alike and, importantly, their millennia of natural history. **References** Addadi, L., Joester, D., Nudelman, F. & Weiner, S. 2006. Mollusc shell formation: A source of new concepts for understanding biomineralization processes. _Chemistry - A European Journal_ , **12** : 980–987. Aguilera, F., McDougall, C. & Degnan, B.M. 2017. Co-option and de novo gene evolution underlie molluscan shell diversity. _Molecular Biology and Evolution_ , **34** : 779–792. Arivalagan, J., Marie, B., Chiappetta, G., Vinh, J., Gallet, X., Lebon, M., M'Zoudi, S., Dubois, P., Berland, S. & Marie, A. 2020. Deciphering shell proteome within different Baltic populations of mytilid mussels illustrates important local variability and potential consequences in the context of changing marine conditions. _Science of the Total Environment_ , **745** : 140878. Bitter, M.C., Kapsenberg, L., Gattuso, J.P. & Pfister, C.A. 2019. Standing genetic variation fuels rapid adaptation to ocean acidification. _Nature Communications_ , **10** : 5821. Byrne, M. & Fitzer, S. 2019. The impact of environmental acidification on the microstructure and mechanical integrity of marine invertebrate skeletons _. Conservation Physiology_ , **7** : coz062. Chan, V.B.S., Johnstone, M.B., Wheeler, A. P. & Mount, A.S. 2018. Chitin facilitated mineralization in the Eastern oyster. _Frontiers in Marine Science_ , **5** : 347. Checa, A. & Harper, E. 2014. Periostracal mineralization in the gastrochaenid bivalve _Spengleria_ . _Acta Zoologica_ , **95** : 196– 208. 

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Telesca, L., Peck, L.S., Sanders, T., Thyrring, J., Sejr, M.K. & Harper, E.M. 2019. Biomineralization plasticity and environmental heterogeneity predict geographical resilience patterns of foundation species to future change. _Global Change Biology_ , **25** : 4179–4193. 

- Waite, J.H. 1977. Evidence for the mode of sclerotization in a molluscan periostracum. _Comparative Biochemistry and Physiology_ , **58B** : 157–162. 

- Waldbusser, G.G., Brunner, E.L., Haley, B.A., Hales, B., Langdon, C.J. & Prahl, F.G. 2013. A developmental and energetic basis linking larval oyster shell formation to acidification sensitivity. _Geophysical Research Letters_ , **40** : 2171–2176. 

- Weiner, S. & Traub, W. 1980. X-ray diffraction study of the insoluble organic matrix of mollusc shells. _FEBS letters_ , **111** : 311 –316. 

- Weiner, S. & Hood, L. 1975. Soluble protein of the organic matrix of mollusc shells: A potential template for shell formation. _Science_ , **190** : 987–989. 

- Weiner, S., Talmon, Y. & Traub, W. 1983. Electron diffraction of mollusc shell organic matrices and their relationship to the mineral phase. _International Journal of Biological Macromolecules_ , **5** : 325 - 328. 

- Weiss, I.M. & Scho nitzer, V. 2006. The distribution of chitin in larval shells of the bivalve mollusc _Mytilus galloprovincialis_ . _Journal of Structural Biology_ , **153** : 264-277. 

- Xia, J., Yu, Z. & Kong, X. 2009. Identification of seven _Crassostrea_ oysters from the South China Sea using PCR-RFLP analysis. _Journal of Molluscan Studies_ , **75** : 139–146. 

- Zhao, R., Takeuchi, T., Luo, Y.J., Ishikawa, A., Kobayashi, T., Koyanagi, R., Villar-Briones, A., Yamada, L., Sawada, H., Iwanaga, S., Nagai, K., Satoh, N. & Endo, K. 2018. Dual gene repertoires for larval and adult shells reveal molecules essential for molluscan shell formation. _Molecular Biology and Evolution_ , **35** : 2751–2761. 

- Zheng, Z., Jiao, Y., Du, X.,Tian, Q., Wang, Q., Huang, R. & Deng, Y. 2016. Computational prediction of candidate miRNAs and their potential functions in biomineralization in pearl oyster _Pinctada martensii_ . _Saudi Journal of Biological Sciences_ , **23** : 372–378. 

- Zheng, Z., Li, W., Xu, J., Xie, B., Yang, M., Huang, H., Li, H. & Wang, Q. 2020. LncMSEN1, a mantle-specific LncRNA participating in nacre formation and response to polyI:C stimulation in pearl oyster _Pinctada fucata martensii_ . _Fish and Shellfish Immunology_ , **96** : 330–335. 


**Fig. 8. Examples of ecological functions served by bivalve shells such as rock oysters. Oyster reefs create habitats for other species such as gastropods, chitons, crustaceans, and algae. Oyster reefs change the structure of rocky shores and the oysters, therefore, are ecosystem engineers. Location: Lantau Island, Hong Kong.** 

_Editors note_ ~ This review was solicited from Alessia Carini after she presented a paper at the Malacological Forum. I suggested the review because I have a special interest in the subject of conchiolin, since I had analysed the amino acid composition of several species of freshwater gastropods in relation to water hardness for my doctoral thesis, presented in 1973. 




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**Research financially supported by the Malacological Society of London** 

## **Taking advantage of contamination: Molecular analyses of solenogaster midgut contents to determine food sources** 

Ludwig-Maximilians-Universita t Munich Dept. Biology II - Systematic Zoology Großhaderner Str. 2 82152 Planegg- Martinsried Email  franzi.bergmeier@googlemail.com 

In 2018, The Malacological Society of London provided an early  career research grant to Lukas Ostermair to support his master thesis. We are very excited that results of this project are now published as a Correspondence piece in _Current Biology_ . 

In his master’s thesis, Lukas tested a molecular approach to investigate the food sources of Solenogastres, a class of worm -shaped and shell-less molluscs. Traditionally, anthozoan cnidarians were proposed as the main food source of these marine molluscs, as undigested cnidocysts were often found during histological investigations of the midgut (Salvini-Plawen 1981), or based on live feeding observations of large-sized species (e.g. Sasaki and Saito 2005). For this project, gut contents were indirectly sequenced from genomic DNA extracts of almost 200 specimens of deep-sea Solenogastres assigned to more than 60 species, constituting their currently known diversity the deep Northwest Pacific (Bergmeier et al. 2019). We used a universal primer pair targeting the nuclear 28S rRNA region and took advantage of a known amplification problem in solenogaster nuclear genes (i.e. their complex secondary structures usually hamper standard PCRs and instead result in contaminations amplified from gut contents (Okusu and Giribet 2003; Meyer et al. 2010)). 

This indirect sequencing of gut contents allowed us novel insights into the feeding ecology of these molluscs. In short, we discovered more than 26 food sources from four different phyla. In addition to cnidarians and annelids, the studied Solenogastres also prey on nemerteans and bivalve molluscs. The majority of investigated solenogasters species seem to have highly specific prey preferences, feeding only on a single food source. Surprisingly, a generalist feeding strategy was only documented in one species. Cnidarians still constitute the most common food source in Solenogastres, but in addition to anthozoans and hydrozoans, some species also prey on benthic siphonophores or scavenge on jelly-fall. 

Finally, we plotted these food sources onto a phylogeny of the investigated solenogaster species and discussed the findings in an evolutionary context. The experimental procedures of this project are detailed in the Supplemental information of the publication. 

We would like to thank _The Malacological Society_ for funding this student project, which resulted in this study on solenogaster food sources and allowed us to gain a better understanding of what’s on the menu for these enigmatic molluscs. The publication is available from the journal’s homepage through a subscription or we will happily share it via e-mail (franzi.bergmeier@gmail.com): 

Bergmeier FS, Ostermair L, and Jo rger KM (2021). Specialized predation by deep-sea Solenogastres revealed by sequencing of gut contents. _Current Biology_ 31(13): R836-R837, https://doi.org/10.1016/j.cub.2021.05.031. 


**Graphical summary of the research project. Inner blue semicircle with some of the investigated Solenogastres species; outer blue semicircle with some of the sequenced food sources (from left to right: Alcyonaceae (Cnidaria), Monostilifera (Nemertea), Trachymedusae (Cnidaria), Leptothecata (Cnidaria), Nuculanida (Mollusca), Terebillida (Annelida)).** 

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## **References** 

**Bergmeier FS, Haszprunar G, Brandt A, Saito H, Kano Y, Jo rger KM (2019). Of basins, plains, and trenches: Systematics and distribution of Solenogastres (Mollusca, Aplacophora) in the Northwest Pacific.** _**Progress in Oceanography**_ **, 178 : 1-17. Meyer A, Todt C, Mikkelsen NT and B. Lieb (2010). Fast evolving 18S rRNA sequences from Solenogastres (Mollusca) resist standard PCR amplification and give new insights into mollusk substitution rate heterogeneity.** _**BMC evolutionary biology**_ **10 (1): 1-12.** 

**Okusu A and Giribet G (2003). New 18S rRNA sequences from neomenioid aplacophorans and the  possible origin of persistent exogenous contamination.** _**Journal of Molluscan Studies**_ **69 (4): 385-387.** 

**Sasaki T and Saito H (2005). Feeding of** _**Neomenia yamamotoi**_ **Baba, 1975 (Mollusca: Solenogastres) on a sea anemone.** _**Venus**_ **64 : 191–194.** 

**Salvini-Plawen Lv (1981). The molluscan digestive system in evolution.** _**Malacologia**_ **21 : 371-401.** 

## **Author’s note** 

**In 2018, my student Lukas Ostermair received a junior research grant for his master’s thesis. We now have finally published the results as a Correspondence piece in** _**Current Biology**_ **. I would like to thank the Society again for their support, especially in relation to young students.** 


**Mussels under threat: combined impacts of ocean acidification and pharmaceutical contamination on** _**Mytilus edulis**_ 

Department of Biosciences, University of Exeter, Stocker Road, Exeter, EX4 4LD, UK * Email: aw429@exeter.ac.uk 

Ocean acidification, the “evil twin” of global warming, is known to be a threat to calcifying molluscs (Gazeau _et al.,_ 2013). Increasing atmospheric carbon dioxide is absorbed by the oceans, decreasing seawater pH and reducing the bioavailability of carbonate ions which are key for calcifiers to build shells and skeletons. In addition, the changes in carbonate chemistry induced by ocean acidification have the potential to change the chemical behaviour of marine contaminants. The associated reduction in seawater pH can alter the bioavailability and toxicity of ionisable organic contaminants to marine organisms. Species which lack the ability to maintain their extracellular acid-base status in ocean acidification conditions, such as the blue mussel _Mytilus edulis_ , may be particularly at risk from this; their internal carbonate chemistry mirrors that of the surrounding seawater, and therefore chemical changes to pollutants could also take place inside their bodies, resulting in altered toxicity effects (Lewis _et al_ ., 2016). 

Research into these potential interactions between ocean acidification and contaminant toxicity for mussels and other molluscs is lacking, despite their potential susceptibility. I designed an experiment to test the hypothesis that the limited ability of _M. edulis_ to regulate its acid-base status in response to ocean acidification will cause predictable changes to the toxicity of contami- 


**Alice Wilson-McNeal conducting some muddy fieldwork in the Exe Estuary, UK.** 

nants. In the experiment, individual mussels were exposed to one of two pharmaceutical drugs at present-day seawater pH and carbon dioxide concentrations, or at simulated pH and carbon dioxide concentrations predicted to be present in the oceans by the year 2100 according to the Intergovernmental Panel on Climate Change. Based on their chemical behaviour, we predicted the toxicity of one of these drugs should increase in this ocean acidification scenario, whereas the toxicity of the other should decrease. We measured haemolymph carbonate chemistry to measure the strength of the mussels’ response to ocean acidification, metabolic rate and ammonia excretion rates to investigate energetic effects, and antioxidant activity and oxidative stress as measures of drug toxicity. The combination of these endpoints would give us an overall picture of the mussels’ general health status as well as the specific effects of the drugs in ocean acidification conditions. 

The experiment was set up in March 2020 but we couldn't access our labs or wild mussels due to the covid pandemic. The Malacological Society graciously 

_**Mytilus edulis**_ **in experimental tanks.** 

offered to extend the duration of the grant, and we were able to postpone the work until restrictions were eased. In the meantime, we marine biologists had to get creative with working from home. I took the opportunity to read more literature than at any other time during my PhD, and came across an additional assay for when we got back into the labs. This assay would give us a greater understanding of the neurophysiological effects of the pharmaceutical drugs. 

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Once we were able to get back into the lab, socially distanced of course, it was a race against time to have the work complet- 

ed before the mussels’ spawning season as this would interfere with their normal physiology. We collected almost one hundred mussels from the Exe estuary and transported them back to the Exeter University aquarium facility. After some fine-tuning of carbon dioxide concentrations in the tanks and confirmation of the experimental doses of the drugs, mussels were finally added to their tanks in September 2020 – six months later than planned. After two weeks we were able to explore the effects of each of the six treatments to the mussels. Whilst changes in acid-base physiology caused by ocean acidification  appeared to affect the toxicity of the two drugs, this was not as clear-cut as we had expected. Some parameters were affected far more than others, and the drug which was expected to become less toxic in ocean acidification conditions had greater effects on some endpoints.  The mussels’ physiological responses to ocean acidification probably influenced the toxicity of the contaminants, but the exact mechanisms are not fully understood. 


**Experimental tanks aerated with an air/CO2 mix to simulate end-of-century ocean acidification conditions.** 


We are indebted to the Malacological Society of London for providing an Early Career Research Grant to conduct this research. This enabled the purchase of additional chemicals to run liquid chromatography-gas spectroscopy analysis of seawater chemistry to validate the findings and additional equipment for experimental work using carbon dioxide. Thank you to my supervisors Dr Ceri Lewis and Dr Rod Wilson for their guidance in writing the grant proposal and designing the experiment, and to Katherine Colvin and Will Davison for their assistance with field and laboratory work. This work was also supported by the University of Exeter Vice Chancellor’s Scholarship for Postgraduate Research. We hope to publish the results from this research this year. 

**Measuring ecotoxicological parameters in the lab.** 


**Using shotgun sequencing for disentangling a taxonomic jumble: the case study of the skeneimorphs Samuel Abalde** Email saabalde@gmail.com 

## **Introduction** 

The Gastropoda comprise the most diverse class of Mollusca, including between 40,000 to 150,000 species adapted to live in all environments. This astonishing diversity has hindered a comprehensive understanding of their evolutionary history, leading to multiple alternative classifications [1] and the subsequent creation of taxonomical categories that do not represent natural groups. 

The family Skeneidae Clark W., 1851 represents an extreme example of such taxonomic conundrum. This family traditionally includes hundreds of species in dozens of genera that are characterized by a minute shell (usually 0.8–2 mm) without nacre, a rhipidoglossate radula and a multispiral operculum [2]. However, more detailed morphological and molecular studies have proved their polyphyletic nature and  the name “skeneimorphs”  was claimed to be more accurate for these snails [3]. For instance, although the family Skeneidae has been assigned to the superfamily Trochoidea [4], some specimens should be considered members of the superfamily Seguenzioidea [3], and there are even cases of former skeneimorphs that belong to the subclass Heterobranchia [5]. 

Molecular studies could help us overcoming this situation. In particular, mitochondrial genomes have been widely used to infer the evolutionary history of different gastropods, and to define a robust classification of these groups [e.g. 6]. In the summer of 2017, I visited the lab of Professor Yasunori Kano (University of Tokyo) aiming to amplify the complete mitochondrial genomes from several species of skeneimorphs from his collection. However, the low amount of DNA (due to the size of these snails), the lack of specific primers (given their polyphyletic origin) and the quality of the probably fragmented DNA, hindered my approach. The main goal of this project was to overcome these limitations by establishing a new methodological pipeline, sequencing low coverage genomes from three skeneimorph species. This would be instead of using the traditional PCR-based approach. 

**Figure 1 – Annotated mitochondrial genome of** _**Leuchorhynchia tricarinata**_ **. Diagram generated with Geneious.** 

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## **Materials and Methods** 

Three species were selected: _Dillwynella vitrea, Leucorhynchia tricarinata_ , and _Protolira thorvaldssoni_ , collected between 2003 and 2012 and preserved in 100% ethanol. Total genomic DNA was extracted from whole specimens using the Qiagen DNeasy Blood & Tissue Kit. Samples were then sent to Macrogen, where an indexed library per sample was constructed using the TruSeq Nano DNA Kit and sequenced in an Illumina NovaSeq platform with a sequencing depth of 30GB. Raw reads were corrected for potential sequencing errors using Rcorrector [7], adapters were removed using Trimmomatic, and reads were quality trimmed using Prinseq v.0.20.4 [8]. Mitochondrial genomes were assembled using NOVOPlasty v.4.2 [9] and annotated by comparison with other Vetigastropoda genomes. Additionally, in order to assess the minimum sequencing depth required to recover complete mitochondrial genomes, reads were randomly sampled to generate 2, 5, 10, 15, and 20GB files. All reads were mapped to the assembled mitochondrial genomes using Bowtie2 v.2.2.6 [10] to separate the nuclear reads, which were then assembled using SPAdes v.3.14.1 [11]. Genome completeness was assessed using BUSCO v.3.0.2 [12]. 

## **Results and Discussion** 

Mitochondrial genomes have been widely used for inferring phylogenetic relationships within Gastropoda, and have helped in the definition of several trochoidean families [6, 13]. Based on this experience, mitogenomes were chosen to study the polyphyletic Skeneidae and identify those species that could belong to different families, but the traditional PCR-based approach was largely unsuccessful. Genome skimming techniques allow the assembly of complete or nearly-complete mitochondrial genomes and the annotation of nuclear markers just from DNA extractions, and it has been successfully used in other contentious groups [14]. 

Between 144 and 211 million reads were generated per species. The complete mitochondrial genome from _L. tricarinata_ (Fig. 1) and _P. thorvaldssoni_ were recovered. The mitogenome of _D. vitrea_ , however, was not circularized. Instead, two fragments of 14,400 and 3,300 bp were assembled. Successive attempts improved this result, although a small fragment between the _trnD_ and the _trnS_ genes was still missing. During the quality control of the extraction, the Macrogen lab team detected a high degradation in this sample, which could explain this result. All mitochondrial genes could be annotated however, proving the reliability of this approach for assembling and annotating the (nearly) complete mitochondrial genome of these minute skeneimorphs. 

Aiming to identify the minimum sequencing depth required the assembling of a complete skeneimorph mitogenome. Different depths were simulated by randomly sampling 6, 16, 33, 50 and 67 million reads for each species. For _L. tricarinata_ 


**Figure 2 – Sequence lengths of the mitogenomes assembled at different sequencing depths for the three species:** _**Dillwynella vitrea**_ **(blue),** _**Leuchorhynchia tricarinata**_ **(orange), and** _**Protolira thorvaldssoni**_ **(gray).** 

and _P. thorvaldssoni ,_ virtually no differences were found among attempts, as the complete mitochondrial genome was recovered in all cases. Small variations in length were detected however, due to the differences in the assembly of the repetitive control region (Fig. 2). 

By contrast, _D. vitrea_ was more troublesome, as at least 10GB were necessary to replicate the results of the full dataset. Based on this experience, DNA quality (or integrity) seems to be an important factor for recovering a complete mitochondrial genome, although this limitation can be bypassed by increasing the sequencing effort [15]. 

Although most of this work was focused on mitogenomes, a big portion of the nuclear genome was also assembled (Table 1), representing between one third and a half of the vetigastropod _Haliotis rufescens_ genome [16]. All three genomes were highly complete, with more than 65% of the metazoan BUSCO genes present in all cases. As expected for a genome assembled from short reads, they are highly fragmented, with the highest N50 of the three in 13,923bp, as reported for other species [e.g. 17]. 

**Table 1: Basic statistics of the assembled nuclear ge-** 

|**Total length (bp)**<br>**Number of contgs**<br>**Largest contg (bp)**<br>**N50**<br>**BUSCO(C/P/M)* **|**_D. vitrea_**<br>**_L. tricarinata_**<br>**_P. thorvaldssoni_**|**_D. vitrea_**<br>**_L. tricarinata_**<br>**_P. thorvaldssoni_**|**_D. vitrea_**<br>**_L. tricarinata_**<br>**_P. thorvaldssoni_**|
|---|---|---|---|
||838,389,779|<br>627,445,078|<br>511,601,414|
||737.371|<br>264,380|<br>376.317|
||148.918|<br>367.191|<br>232,540|
||3.154|<br>13.923|<br>2.873|
||35.9/30.6/33.5|<br>82.2/12.5/5.3|<br>44.7/31.6/23.7|



*Percentage of C = Complete; P = Partial; M = Missing genes 

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## **Future work** 

This project was conceived as a proof of concept to evaluate the performance of genome skimming for studying skeneimorph gastropods. After these successful results, this protocol will be now extensively applied to other skeneimorph samples, aiming to identify highly divergent lineages that might represent independent families. Thus far, one of the goals included in the proposal was inferring the phylogenetic position of the Skeneidae within Trochoidea. However, the inferred tree topology was highly variable depending on the analysis. This project will surely benefit from an increased taxon sampling. 

## **References** 

1. Aktipis, S.W., _et al_ . 2008. Gastropoda: An overview and analysis. In: _Phylogeny and Evolution of the Mollusca_ , pp. 201-237, edited by Ponder, W.F. and Lindberg, D.R. University of California Press, Berkeley. 

2. Haszprunar, G., _et al_ . 2016. Towards a sound definition of Skeneidae (Mollusca, Vetigastropoda): 3D interactive anatomy of the type species, _Skenea serpuloides_ (Montagu, 1808) and comments on related taxa. _Organisms Diversity & Evolution,_ **16** : 577-595. 

3. Kano, Y., _et al_ . 2009. Morphological, ecological and molecular characterization of the enigmatic planispiral snail genus _Adeuomphalus_ (Vetigastropoda: Seguenzioidea). _Journal of Molluscan Studies_ , **75** : 397-418. 

4. Williams, S.T. 2012. Advances in molecular systematics of the vetigastropod superfamily Trochoidea _. Zoologica Scripta_ , **41** : 571-595. 

5. Ware n, A., _et al_ . 1993. Systematic position of three European heterobranchian gastropods. _The Veliger_ , **36** : 1-15. 

6. Uribe, J.E., _et al_ . 2017. Denser mitogenomic sampling improves resolution of the phylogeny of the superfamily Trochoidea (Gastropoda: Vetigastropoda). _Journal of Molluscan Studies_ **, 83** : 111-118. 

7. Song, Li, et al. 2015. Rcorrector: efficient and accurate error correction for Illumina RNA-seq reads. _GigaScience_ **, 4** (1): s13742-015. 

8. Schmieder, R., _et al_ . 2011. Quality control and preprocessing ofmetagenomic datasets. _Bioinformatics,_ **27** : 863–864. 

9. Dierckxsens, et al. 2017. NOVOPlasty: de novo assembly of organelle genomes from whole genome data. _Nucleic acids research,_ **45** (4): e18-e18. 

10. Langmead, B., _et al_ . 2012. Fast gapped-read alignment with Bowtie 2. _Nature Methods_ , **9** : 357-359. 

11. Bankevich, A., _et al_ . 2012. SPAdes: a new genome assembly algorithm and its applications to single-cell sequencing. _Journal of Computational Biology_ , **19** : 455-477. 

12. Sima o, F.A., _et al_ . 2015. BUSCO: assessing genome assembly and annotation completeness with single-copy orthologs. _Bioinformatics_ , **31** (19): 3210–3212. 

13. Guo, E., _et al_ . 2021. Mitogenomic phylogeny of Trochoidea (Gastropoda: Vetigastropoda): New insights from increased complete genomes. _Zoologica Scripta_ , **50** (1): 43-57. 

14. Sanchez, G., _et al_ . 2021. Phylogenomics illuminates the evolution of bobtail and bottletail squid (order Sepiolida). _Communications Biology_ , **4** (1): 1-9. 

15. Alsos, I. G., _et al._ (2020). The treasure vault can be opened: Large-scale genome skimming works well using herbarium and silica gel dried material. _Plants,_ **9** (4): 432. 

16. Masonbrink, R.E., _et al._ (2019). An annotated genome for _Haliotis rufescens_ (Red Abalone) and resequenced green, pink, pinto, black, and white Abalone species. _Genome Biology and Evolution,_ **11** (2): 431–438. 

17. Barghi, N., _et al._ (2016). Structural features of conopeptide genes inferred from partial sequences of the _Conus tribblei_ genome. _Molecular Genetics and Genomics_ , **291** (1): 411-422. 




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## **Book reviews** 

Published  by the Royal Society  in special issue 1825 of volume 376 

Print ISSN:0962-8436     Online ISSN:1471-2970 

Genome sequencing seems now so commonplace, a critical part of the Covid-19 response, embedded in personal healthcare, and the reporting of genome sequencing of ‘newsworthy’ organisms, that it would be understandable if the general public held the misperception that genome sequencing was trivial, with the science at a stage where any organism could have its entire genome sequenced quickly, cheaply, with apparently little effort. It is preaching to the converted to argue that molluscs display a wide variety of phenotypes of interest addressable through genomic methods. Why then are, particularly high-quality molluscan genomes, under-represented in genome databases, how can this be changed, and what even can we learn from molluscan genomes? This special issue of _Philosophical Transactions of the Royal Society_ : ‘Molluscan genomics: broad insights into future directions for a neglected phylum’ reports on a meeting, ‘Pearls of wisdom: synergising leadership and expertise in molluscan genomics’ held in 2019 and considers, in a series of 14 papers, particular aspects of, or challenges to, molluscan genomics. From technical difficulties in extracting the high molecular weight DNA necessary for genome sequencing (molluscs seem to have evolved a number of characteristics which make this extremely difficult), right through to front-end database availability of genomes accessible by all, and from the current knowledge of epigenetic modification of genomes through to the state of play of molluscan phylogenomics. The introductory paper to the volume by Angus Davidson (University of Nottingham) and Maurine Neiman (University of Iowa), the meeting conveners, sets the scene for these diverse papers, explaining the current state of play of molluscan genomics and the (quirky) history and rationale for this conference. 

Adema reviews difficulties with extracting high quality molluscan DNA, something anyone studying molluscan genetics will be abundantly aware of. This article clearly presents the issues affecting extraction e.g. mucopolysaccharides and pigments, and difficulties with qualitative assessment of molluscan genomic DNA, an article I am sure will be extremely useful to all undertaking such procedures. 

We sit at an exciting juncture where not only are large national (e.g. Darwin Tree of Life project) and international efforts underway to sequence the genomes of all species but individual labs can feasibly sequence whole genomes. In recent years the Oxford Nanopore Technologies (ONT) sequencer, a device that in its simplest form is a small USB powered sequencer, has been demonstrated to be deployable anywhere from field diagnostic laboratories in Liberia to the International Space Station. However, the latest iterations of this and the bioinformatics methodology to deal with these data make it usable in any laboratory for genome sequencing. Sun _et al._ take molluscan ONT data and benchmark a range of assembly pipelines offering advice for the optimum methods dependent on the size and heterozygosity of your molluscan genome. 

To be useful, molluscan genomes must be accessible, not just as raw data available only to those with the necessary bioinformatic skills, but through freely accessible online genome databases. Caurcel _et al._ present MolluscDB (https:// molluscdb.org/) an access route for both currently available and future mollusc genome assemblies. Whole genomes represent a rich dataset for phylogenetics and molluscan phylogenetics (with  contentious evolutionary relationships, particularly within classes) may benefit greatly from the large datasets genomics can provide. Sigwart _et al._ consider where research activities should be directed in future years to optimise the utility of these, noting not that although most molluscan diversity remains unsampled, current genome sequences are typically from recently derived taxa and future efforts must concentrate on early diverging branches and groups with poor sampling (e.g. Aplacophora, Schaphopoda, Polyplacophora). 

Whole-genome sequencing data opens up a swathe of opportunities for understanding the driving forces of speciation. Chueca _et al._ use data from two species of the terrestrial pulmonate _Candidula_ to dissect the role, and extent of gene flow, to understand ecological speciation in these snails. 

Whole mitochondrial genomes have been attainable for longer than whole (nuclear) genomes, although they now can be extracted from whole genome data. Ghiselli _et al._ review the current knowledge on the gene content and organisation of molluscan mtDNA and the utility of these sequences in a phylogenetic context. 

The term ‘model organism’ is widely used in biology. In their paper, Davison and Neiman discuss why there are no molluscan model organisms and what criteria must be met for any mollusc to be regarded as such, including the genomic and genetic resources necessary. 

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Other papers are perhaps more relevant to those interested in molluscan molecular genetics. Rosani _et al._ review current knowledge concerning molluscan miRNAs (non-coding regulators of gene expression) and Ma nner _et al._ consider a potential epigenetic regulator of gene expression, methylation, inferring methylation patterns across 140 molluscan species. Calcino _et al._ show that molluscan genomes not only show evidence of hemizygous regions (with large chromosomal sections differing in presence/absence) but that these regions typically contain genes involved in transposition, DNA repair and stress response. Other structural variation is detectable in oyster ( _Crassostrea virginica_ ) genomes (Modak _et al._ ). These duplications contain either genes or exons and are hypothesised to underpin this species’ adaptive and evolutionary success. McCartney utilises genomics and classical genetics to locate and study the genes involved in byssus production from a _Dreissena polymorpha_ genome assembly, comparing byssus proteins with those from quagga and blue mussels. Molluscs of aquacultural importance are perhaps over-represented in the (limited) available genomes. Potts _et al_ . discuss how genomic data and technologies may aid in improvements to disease resistance including how genomic data can be interrogated for suitable loci for CRISPR modification. Finally, Seppa la _et al_ . discuss the adaptive evolution of the gastropod immune response, reviewing the studies on this and discussing how genomic, or more particularly transcriptomic data may provide insights into this. 


This volume is therefore not just a collection of papers detailing molluscan genomes, but a state-of-the-art summary, offering suggestions, both technical and reflective for how to move forward. Whilst it will, unsurprisingly, be likely of most interest to those directly involved in genomics (and not just of molluscs), it contains a variety of papers of interest to many broad molluscan interests, and demonstrates the utility of genomic data to throw light on important questions in molluscan biology. 


Dr Craig Wilding Reader in Evolutionary Genetics Liverpool John Moores University, L3 3AF 


**----- Start of picture text -----**<br>
Book and film<br>Other minds; the Octopus and the evolution of intelligent minds<br>     by Peter Godfrey Smith  (2018).  Pubished by William Collins, London   ISBN: 978-0-00-822629-9<br>BBC iPlayer - Natural World - 2019-2020: The octopus in my house<br>       Presented by Professor David Scheel<br>**----- End of picture text -----**<br>


Peter Godfrey Smith (PGS) begins the book by describing ‘Octopolis’, an octopus ‘city’ over which he scuba dives on a shallow sea floor on the east coast of Australia. Octopolis is  a habitat to which the author returns several times during the narrative. It is a rather strange biotope, in that octopods are not usually social animals. 

The author takes an evolutionary approach to the question of how sentience, intelligence and consciousness fit into the physical world.  As a philosopher, he wants to know how it feels to be an animal such as a cephalopod – “If we want to understand other minds, the minds of cephalopods are the most other of all” emphasising that “this is a philosophy book as well as a book about animals and evolution”. 

Early in his narrative, PGS describes what is meant by ‘seeing’ for a bacterium, including both the seeing its environment and seeing its bacterial colleagues. The relevance of this to malacology is that, for example Hawaiian squid host bacteria in their body. The bacteria produce light through a quorum-sensing mechanism, and the host squid incorporates the process into its development of camouflage. 

Two possible narratives for the development of multicellularity are discussed, comprising sponges and comb jellies. The author observes that Ediacarans in the Precambrian beds near Adelaide may have had little to do with each other “munching on the mat (of) filtered food from the water … hardly interacting at all”, since there is little evidence of predation. It seems that at this time, the nervous system was used for coordination rather than the sensori-motor control as used for, for example, in the processes of defence or attack. PGS then nicely obfuscates his own argument by raising the issue of jellyfish stings. Who was being stung, and why? 

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## >CONTINUED 

The Cambrian metazoan explosion is described as a process of animals being “involved in each other’s lives”. Through natural selection and evolution this leads to the production of eyes, antenna and claws, The book is peppered with nuggets of biology - for example box jellyfish can navigate, moving at three knots, by watching external landmarks on the seashore. Eventually, complex, active bodies arise in only three taxa, these being the chordates, arthropods and cephalopods. The writing is both poetic and enigmatic in places, for example “until in one example of this sensory and behavioural entangling, a rubber encased mammal and a color changing Cephalopod find themselves staring at each other in the Pacific ocean”  PGS moves on into the evolution of molluscs, pausing on _Nautilus_ as a lone survivor with 200 million years of heritage. 

Homing in on the Cephalopods, PGS observes that they evolved to “smart”. He examines the mismatch between experiment and anecdote as a familiar problem in animal psychology, but it is particularly relevant to octopus behaviour. He describes experiments and observations on the extraordinary behaviours of cephalopods in the open sea, in Octopolis and in the laboratory. 

Mammal brains can be mapped but it is difficult when the octopus has most of its neurons in its arms. Because of recurrent neural connections, octopuses have a form of short term memory in the arms. Nevertheless, in experiments, the octopus uses its eyes to guide its arms through a maze to a reward.  The arms may also have sufficient independence to do some local exploration, self-motivated by the arms. PGS asks why an octopus has as many as 0.5 billion neurons and investigates ideas of primate behaviour developed by Katherine Gibson in the 1980s.  Gibson firstly identified food gathering that requires little manipulation, for example a frog catching an insect.  Secondly there is extracted foraging where choices are made within the context, for example removing food from protective shells. In relation to this, PGS reconsiders the evolution of nervous systems, contrasting sense or in motor views with action-shaping views, suggesting that the first nervous systems were primarily for coordination within the organism.  He suggests the octopuses central nervous system imposes a top down order but also allows turned each arm to be an intermediate scale actor.  Thus, the octopus has a kind of mixed (nervous) economy.  The outcome is that it is not clear to us where the brain ends in an octopus, since the nervous system runs throughout a body which has “no parts, an old joints, no natural angles”. 

What it feels like to be another organism in its world is then considered. He investigates the ideas of consciousness, discussing issues such as perceptual constancy (how an object remains recognisable of the same object as the viewpoint changes) and nervous integration in a range of taxa, concluding that brain activity which leads to visual experience causes the building of an inner model of the external world in the nervous system.  In exploring consciousness, the author takes the example of pain, noting that some animals don’t seem to feel pain.  For example, they don’t groom or protect an injured part of the body but just continue with whatever they were doing.  He points out that subjective experience comes from the modulation of the nervous system, that is, from registering things that matter and not just merely running the nervous system. In developing this argument, PGS invokes the work of Dehaene, Barrs, Millner and Goodale, seeing consciousness as a useful term for subjective experience that is “unified and coherent in various ways”.  He takes the special case of the octopus, describing a range of experiments which  reinforce the idea that octopuses have delegated a certain amount of local control to their arms. At this point, PGS again wonders what it would be like to be an octopus. For the octopus nervous system, PGS accepts the metaphor that it may resemble jazz improvisation, where the final result emerge as out of a continued give and take of musical information between the players. 

The colour changes of the giant cuttlefish is next for consideration. PGS is puzzled by the metabolic cost of the immensely expressive colour changes in an animal that apparently does not have colour perception, and in relation to cephalopod camouflage asks “How can you match colors you cannot see?”  The relationship between colours and behaviour of shapes is explored; for example, when are a cuttlefish holds its arms up like horns, in an aggressive gesture.  Many of these engaging examples of relationships between the observer and the Cephalopod take place in Octopolis and PGS describes examples of the animals exhibiting blatant curiosity, with associated colour changes.  As a general biologist, I was pleased to see that PGS includes an analysis of the mechanisms of these colour changes. The descriptions in the book are eloquent but to really assimilate what is happening, I recommend the  BBC video available at BBC iPlayer - Natural World - 2019-2020: The Octopus in My House Here, David Scheel  Professor at Alaska Pacific University spends a year in a personal relationship with an octopus in an Octopolis in the USA. Finally in relation to colour, PGS concludes that colour changes in the giant cuttlefish are an inadvertent expression of the animals inner processes; he offers colour plates in evidence and again, his thesis is efficiently supported by the BBC film documentary. 

The book now enters a more philosophical vein, passing from Hume to Vygotsky via Darwin. The relationships between speech and thinking in humans is explored by reference to the psychologists’ use of the concepts of System I (thinking uses habits and intuitions) and System II (which is slower, users proper rules of reasoning, and tries to look at things from several angles, being “ponderous and powerful”). At this point, I notice a comment I have written on a page in the book “ _has he abandoned the cephalopods now?”_ but this is not a negative criticism. PGS can’t stop being a philosopher and so, he makes an excursion into thinking about one’s own thoughts. As this stimulating chapter ends, the author brings his readers back to the molluscs. 

The short life of a cephalopod allows PGS to start wondering why anything gets old and dies, and he discusses the possibility that the death of aging individuals saves resources for the young, thereby benefitting the species. The evolutionary principle of mutation-selection balance is examined, including the logic that individuals who live long and without deleterious mutations are likely to have more offspring.  By contrast, everyone will carry some mutations and it will appear to an observers that they have been programmed to decline.  PGS develops his argument by considering semelparous and iteroparous life cycles.  Semelparous organisms have-big bang reproduction, producing offspring in a single short season.  Iteroparous organisms produce several generations over a more extended period.  Female octopuses are an extreme case of semelparity since they die after a single pregnancy. 

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## CONTINUED> 





**----- Start of picture text -----**<br>
  The den<br>**----- End of picture text -----**<br>


**First encounter** 

**Exploration of a relationship** 




**A bitten off arm regrows** 

**A child gets involved As a protection, the octopus forms a ball, armour plated with shells** 




**----- Start of picture text -----**<br>
It is investigated by a predatory fish…...<br>**----- End of picture text -----**<br>



**----- Start of picture text -----**<br>
...and in a blur, rolls away in its armour.<br>**----- End of picture text -----**<br>


**Some still images from The octopus in my house** 

In the final chapter PGS returns to Octopolis, recounting days when he would see more than a dozen octopuses roaming, grappling or just sitting in an area a few metres in diameter. In an observation supported by the BBC documentary, he engagingly describes a Nosferatu pose where a startled octopus colours itself black and raises a dark cloak of mantle forward, over its head.  PGS wonders how Octopolis could originats, and comcudes that it is caused initially by octopuses eating scallops and leaving their shells as debris.  In a positive feedback loop, this debris allows other octopuses to dig out further stable dens so that the environment comprises more and more of the shells themselves, in a case of ecosystem engineering.  All this seems to lead at Octopolis to an unusual social life of high densities and continuous interaction. 

Finally, PGS summarises his argument in relation to the cephalopods and extends it into the negative impacts humans activities have on oceans.  The film also ends on a pensive note, when the main cephalopod protagonist comes to the end of its natural life.  Its death evidently deeply affects its human companion. 

I found the book and the film created a thought-provoking synergy, and I strongly recommend both to anyone who nurtures an interest in molluscs. 

Reviewer - Georges Dussart 




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## **Annual report of Council for 2021/202** 

## **delivered by the President, John Grahame at a Virtual Annual General Meeting** 

## **MEMBERSHIP** – no report 

## **FINANCE for the financial year ending 31[st] December 2020 (reported by Katrin Linse** ). **Significant activities** 

In 2020 the Malacological Society of London had proposed meeting at the Marine Biological Association in Plymouth on "Limpets 2020" that had to be cancelled with a week’s notice due to Covid-19, and the annual Molluscan Forum for Young Scientists was held virtually. The Society published the members' bulletin "The Malacologist" and in cooperation with Oxford University Press the "Journal of Molluscan Studies". 

## **Public benefit** 

The charity's objects are for the public benefit because increasing public knowledge is required as molluscs form an important part of the global biodiversity and ecosystem stability which can have effects on human health and are a human food source. 

## **ACHIEVEMENT AND PERFORMANCE** 

## **Charitable activities** 

In 2020 the main charitable activities were the funding of eleven research projects, of which nine claimed the grants (total cost of £13,342.25), eight to young researchers and malacologists in non-permanent positions and one to senior researchers, the support of 3 students and young researchers to attend conferences and training courses, an activity highly impacted by Covid-19 as the Society usually supports around 20 scientists here. 

## **FINANCIAL REVIEW** 

The finances of the Malacological Society have been pleasing during 2020 with an overall gain of £52263. This gain is explained by a gain in the Fixed Interest and Investment funds and lower awards and meeting expenditure. Our investments had an overall gain of £28,997 (comparing market value at 31 December 2020 with market value at 31 December 2019), with the COIF Investment Fund making a gain of £25,132 and the COIF Fixed Interest Fund a gain of £3,865. During 2020, no funds were transferred from the current account to savings accounts. Separately, the profit-share from the publication of the Journal of Molluscan Studies in 2020 provided the Society with most of its income contributing £49,396. The Editor of the Journal, Dr Dinarzarde Raheem, and the Assistant Editors are to be commended for their hard work contributing to the publication of our scientific journal. In addition, sales of the digital archives provided £506.50 of income. 

In 2020, more funds were used for research awards, being £13,342k in 2020 compared to £8,696k 2019, while travel awards remained similar, but there was reduced spending on Council meeting and Forum travel awards as meetings after January 2020 were virtually. The Society (MSL) spent less money in 2020 compared to 2019, this was mainly based on less expenses paid for meetings and to the JMS editor for attendance expenses Malacological conferences to promote our journal and network with potential assistant editors and authors. 

## **MEETINGS** 

## _**The AGM**_ 

The 127[th] AGM was held as a virtual meeting _via_ Zoom, on the 4[th] of June, the same day as the June Council but as a separate meeting. Very unfortunately it was not only delayed by problems arising from the Covid-19 pandemic, but also was unaccompanied by any malacological presentations because the anticipated Limpets 2020 meeting was necessarily cancelled at very short notice. 

## _**The Molluscan Forum**_ **report-** 

## **ed on by Phil Hollyman** 

The annual molluscan forum was held on the 19th of November 2020, for the first time this meeting was hosted virtually _via_ Zoom. Over 170 people registered in advance for the event, representing about double the delegates of recent inperson meetings. The shift to a virtual platform had a clear impact on accessibility for international delegates, improving the ability of many people to attend and present. 


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## >CONTINUED 

This year also saw the introduction of quick-fire talks in place of poster sessions, giving each speaker five minutes and two slides to present their findings. Overall there were 50 applications for full and quick-fire talks, which were presented during three full and three quick fire sessions throughout the day. **The Oxford prize, awarded annually for the best early career talk, was given to Lauren Eggleton (Sheffield University) for her talk titled: ‘A Sticky Situation: Investigating the Contradictory Nature of Gastropod Mucus’.** 

## **PUBLICATIONS** 

## _**The Malacologist**_ **reported on by Georges Dussart** 

In the 2020-2021 operating year, two issues of The Malacologist were posted to the website. 

The August issue always includes the President’s report of Council, which summarises the activities of the Malacological Society of London over the previous year. Issue 75 (August) of The Malacologist was an extended edition of 53 pages; this pagination largesse was partly due to papers which discussed the future of the Malacological Society of London. There was also an unusually large number of research reports. An article by Robert Cameron on The future of the Society and the Journal of Molluscan Studies in the age of Open (and digital) Access led to articles by Trevor Parry Giles (Crafting a new normalcy) and Robert Harington (Why scholarly societies are vitally important to the academic ecosystem). The research reports included Cryptic and invasive freshwater Galba snails by Pilar Alda, Towards a first molecular phylogeny of Caecidae micro snails by Christina Egger et al., Preliminary data on the biodiversity of solenogasters in the China Sea by M. Carmen Cobo, BrdU-based sperm-labelling protocol in a hermaphroditic freshwater snail species by Yumi Nakadera et al. and Physiological performance of an invasive freshwater gastropod under elevated CO2-induced acidification by Andrew A. David. As always, it was gratifying for the Society to receive a report from an Annual Award winner, in this case Samuel Abalde’s report on Phylogenomics and comparative transcriptomics of West African cone snails. The issue also included a book review and two memoria (John Allen and Ron D’Or).  Finally, the issue included an invited article from Jake Goodall et al. entitled Emerging from the shell: development of the common whelk as a novel model of phenotypic variation. The invitation arose from a series of Forum presentations given on this subject by the Icelandic group. 

The February issue (76) of _The Malacologist_ had, as its core, the abstracts of the Molluscan Forum for 2020. This 41-page issue included two research grant reports. Kiran Liversage et al. reported on Intertidal molluscs on the ecologically dynamic Saurashtra peninsula (Gujarat, India) and the report from Eduardo Sampaio, concerned Disentangling multi-partner effects on collaborative hunting between octopus and fishes. As increasingly happens for research reports, this report was associated with video recordings. The issue included an essay from Council member Robert Cameron which summarised his experience of academic publishing over the past decades; Robert intriguingly entitled the essay Tales from the Palaeolithic. Robert’s essay was prompted by the termination of paper-printing of the Journal of Molluscan Studies (JMS); paper issues are now replaced by digital-only access. This event also prompted the article written by Alan Hodgson, another Council member, From print to on-line, a historical review of the Journal of the Malacological Society of London which reviewed the history and editorship of the JMS since its inception.  A further contribution by Council members in this issue was the position paper from Lauren Sumner-Rooney on  Diversity, Equality and Inclusion in the Malacological Society of London  prompted by the ‘Black Lives Matter’ movement. 

_The Malacologist_ continues to present a useful synoptic view of malacology, especially amongst young researchers. The Malacologist has an ISSN number (ISSN 1759-1406) and is therefore citable, so it is in the interest of awardees to present reports on their Society-funded research or travels. 

## _**The Journal of Molluscan Studies**_ **reported on by Dinarzarde Raheem** 

The ISI impact factor for the _Journal_ in 2019 increased to 1.461 (compared with 1.345 in 2018, 1.483 in 2017, 1.250 in 2016 and 1.185 in 2015). The _Journal_ stands at number 63 in the ISI list of 168 zoological journals (it was 65 out of 170 in the previous year). Our chief competitor, _Malacologia_ , had a remarkably high impact factor of 13.50 in 2019, reflecting the fact that during the two-year citation window there has been just a small number of citable items in this journal, of which one (Bouchet _et al._ ’s 2017 classification of gastropod and monoplacophoran families) has been cited nearly 300 times. The _Journal_ continues to be truly international in terms of the geographical distribution of its authors; for volume 85 (2019) the corresponding authors represented 19 countries (of which the leaders were 16% Germany and 13% Japan). The average publication time from receipt to Advance Access publication was 10.4 weeks for 2019. 

Circulation for the Journal in 2020 was 55 institutional (of which 36 were online-only and 19 print-only) and 156 membership subscriptions (compared with 30 and 157 respectively for 2019). In addition, a further 2,530 institutions have electronic access to the Journal through publishers’ collections (includes migrated figures; compared with 2,348 in 2019). The new pricing structure has been fixed for 2021. The Journal moved to being online only from January 2021, with the cost for an online-only subscription set at £552. Please see https://academic.oup.com/mollus/subscribe for more information. 

Volume 86 (2020) contained 45 papers, research notes and review articles, totalling 434 pages (the preceding volume totalled 452 pages). In total, 127 manuscripts were submitted in 2020 (an increase of 8.5% on the 117 in 2019) and the acceptance rate was 33%. The image of the chiton _Chiton albolineatus_ on the cover of Volume 87 was kindly donated by Douglas J. Eernisse. 

Our board of Associate Editors is now: Coenraad Adema (immunology, genomics, parasitology), Thierry Backeljau (molecular phylogenetics and genetics), Liz Boulding (population and reproductive biology), Robert Cameron (ecology and genetics of terrestrial gastropods), Richard Cook (agricultural malacology, physiology, feeding behaviour), Simon Cragg (life histories, sense organs), Mark Davies (marine ecology and behaviour), Dan Graf (freshwater bivalves), John Grahame (population genetics, morphometrics), Liz Harper (marine bivalves), Gerhard Haszprunar (microanatomy, 3D reconstruction, minor molluscan classes), Bernhard Hausdorf (terrestrial gastropods), Michal Horsa k (ecology and biogeography of 

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## >CONTINUED 

terrestrial gastropods), Yasunori Kano (systematics of vetigastropods, tropical ecology), Joris Koene (reproductive behaviour of gastropods), Manuel Malaquias (opisthobranchs), Peter Marko (marine biogeography and phylogenetics), Pablo Martí n (freshwater ecology, life history), Ellinor Michel (ecology, freshwater gastropods), Jeff Nekola (community ecology of terrestrial gastropods), Nicolas Puillandre (neogastropods), Ellen Strong (freshwater and marine caenogastropods), Janet Voight (cephalopods), Janice Voltzow (microscopic anatomy), Heike Wa gele (opisthobranch biology), Tony Walker (biochemistry, immunology, cytology), Suzanne Williams (molecular phylogenetics and genetics), and Yoichi Yusa (general ecology and behaviour). Nerida Wilson has temporarily stepped down from the editorial board. 

I would like to thank all the members of the editorial board, those members of the international malacological community who have contributed to the review process, the staff of Oxford University Press, particularly Cailin Deery and Jan Webster, and the production teams at Aptara Incorporated (led by Akash Mahajan) and SPi Ltd, Philippines (led initially by Mackie Fernandez and latterly by Ronel Mirano and Erica Fajardo), for their work on behalf of the _Journal_ . 

## **WEBSITE reported on by John Grahame and Tom White** 

- Stefan Senk (http://www.senktec.com/stefan senk/ ), who built the current website, is in the process of making some much needed revision and overhaul of it. John Grahame has started up the learning curve of editing the site, to bring its content up to date. This is very much a work in progress. 

## **FACEBOOK & TWITTER reported by Chong Chen and Lauren Sumner-Rooney** 

The Facebook page continues to be a useful and well-used part of our Web presence. It has proved particularly valuable in communicating information the Society wishes to publicise quickly. 

This year we have also set up a Twitter account to enhance communication. This has also proved successful – set up in July 2020, it has attracted 383 followers with 64.4K views of Tweets. 

If you have suggestions / comments on these pages, please do not hesitate to contact Chong Chen, _Facebook Manager_ or Lauren Sumner-Rooney, _Twitter Manager_ . 

## **AWARDS (reported by Jonathan Ablett)** 

Overall, the Society is very pleased with the number of applications that it receives for Travel Awards and Research Grants however the number of applications received for Travel Awards was severely reduced due to the current global health situation. The schemes seem to be achieving their global aim to enable young scientists to engage in malacological research activity both in the laboratory/field and at meetings. Citable reports from researchers, funded through both schemes, appear in _The Malacologist_ . 

The Society aims to make the following awards annually. 

Travel Awards - at least 5 each of up to £500 for Society members, £300 for non-members 

Research Grants - at least 5 each of up to £1500 

Application forms and guidance notes for both schemes have been updated recently and can be downloaded from The Society’s website. 

## **Travel Awards** 

In 2020 there were 2 rounds of Travel Awards, June and December. Due to the global Coronavirus pandemic the Society only received 1 application for awards to travel and was able to fund this request. All Travel Award applications are reviewed by an Awards Committee. The Society is pleased to have announced the following awards. 

## _June Travel Awards_ 

No applications received and no awards made. 

## _December Travel Awards_ 

Hannah Lee (Heriot-Watt University) **£300** for attendance at ‘NORA 4 (Native Oyster Restoration Alliance)’ Middelburg (The Netherlands), 4th – 6th May 2021 

A total of **£300** was allocated by The Society for Travel Awards. All applicants have been notified of the outcome. Note that this amount does not necessarily reflect actual ‘spend’ as occasionally students withdraw from the intended visit. 

## **Research Grants for 2020 - Senior Research Grants & Early Career Research Grants** 

The society award both Senior Research Grants and Early Career Research Grant. Early Career Research Grants are conferred on students and researchers without professional positions, but without regard to nationality or membership of The Society. Senior Research Awards are aimed at established researchers in professional positions, but without regard to nationality. Applicants for Senior Research Awards must be members of The Society. Early Career Research Grants will be reviewed by MSL council members and Senior Research Grants will be reviewed by a Reviewers Panel including both council and non-council members invited for that purpose. 

4 

## **Early Career Research Grants** 

By the closing date of 15[th] December 2020, the Society had received 10 applications from workers from 10 institutions in 6 different countries. In general, the scientific quality of the research projects submitted was excellent. 

On behalf of the Society, I would like to formally thank the members of the Grants Review Panel for their hard work in reviewing all applications. The Panel has agreed the following awards, in alphabetical order. 

- A. Bhosale (Foundation for Biodiversity Conservation, India), **£1500** 

   - ‘A survey of the land-snail fauna of the northern Western Ghats, India’ 


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## >CONTINUED 

- A. Buckner (University of Lincoln, UK), **£1255** 

   - ‘Immnological respond of _Glaba truncatula_ when infected with _Fasciola hepatica’_ 

- C. Drerup (Cambridge University, UK), **£1500** 

   - ‘Adaptions to visually challenging environments in cuttlefish’ 

- H. Lipae (University of the Philippines Los Ban os, Philippines), **£1500** 

   - ‘Diversity and Community Assembly of Camaenid Land Snails in Karsts on Luzon Island’ 

- L. Martin (Institute of Marine Sciences, University of Portsmouth, UK), **£1500** 

   - ‘Laboratory testing of the efficacy of novel furfurylation treatments and naturally durable, tropical hardwood species against settlement by larvae of the shipworm, _Lyrodus pedicellatus_ and other teredinid molluscs.’ 

- O. Ojeda (University of Ma laga, Spain), **£1250** 

   - ‘Molluscan fauna of mud volcanoes of the north-eastern Gulf of Ca diz: biodiversity and eco-biological aspects.’ 

- K. Zając (Jagiellonian University, Poland), **£1500** 

   - ‘Removing taxonomic obstacles in the genus _Deroceras_ by means of integrative redescriptions of selected taxa.’ 

Therefore **7** Research Grants have been funded at a total cost of **£10,005.** The success rate was **70%** . The Grants Review Panel would like to emphasise that the quality of all applications was high and that it funded as many excellent projects as possible. Applicants have already been formally notified of the outcome of their application. 

## **Senior Research Grants** 

By the closing date of 15[th] June 2020, the Society had received only 1 application for the Senior Research Grants. On behalf of the Society, I would again like to formally thank the members of the Grants Review Panel for their hard work in reviewing all applications. The Panel has agreed the following award: 

- Dr Andrew David (Clarkson University, USA) 

   - **£923.42** ‘Testing the efficacy of a popular synthetic molluscicide (Niclosamide monohydrate) for controlling invasive mystery snails (Mollusca: Viviparidae) in North America). 

## **The Annual Award** 

## The Society received no nominations for the 2020 Annual Award. 

## **The Oxford Prize for Malacology** 

The Oxford Prize for Malacology is awarded annually for the best presentation at the Molluscan Forum, is generously supported by Oxford University Press, publisher of the Society’s journal. The 2020 winner is Lauren Eggleton (University of Sheffield, UK) for her talk entitled ‘A Sticky Situation: Investigating the Contradictory Nature of Gastropod Mucus.’ 

## **OFFICERS & COUNCIL** 

This has been my third and final year as President, and it has brought unique challenges. Council met on 22[nd] January 2020 in the Natural History Museum, aware of reports of a new respiratory virus which had emerged in China but not particularly perturbed by them. What may now seem like complacency would be short-lived. The lead up to our proposed AGM in the context of the proposed Limpets 2020 meeting became a time of increasing anxiety, culminating in the cancellation of the Limpets meeting and the postponement of our AGM. This eventually took place in June (see above) using the Zoom platform, a novelty for many though by no means all of us. One of the discussions was about the Forum, and although the pandemic situation appeared to be improving in June it seemed foolhardy to contemplate a physical Forum symposium for November. Therefore the Forum was moved to Zoom, and proved to be an outstanding success with a stronger international participation than ever (again, see above). Council meetings in June and October 2020 and January 2021 have been in Zoom, we have yet to see how things work in June 2021, and later. While we all long to meet physically again, the Zoom platform has allowed much easier access to meetings for geographically distant councillors, and this has been welcome. Our _Journal_ has moved to online only – a change which would have happened in the not too distant future anyway, but which was brought forward in response to pandemic difficulties. It’s in good health and in the hands of an excellent Editor in Chief, Dinarzarde Raheem, supported by a team of Associate Editors. _The Malacologist_ with Georges Dussart as editor continues to function as our interesting and wonderfully illustrated newsletter and reporter of the malacology we have supported through Awards. This past 12 months it has documented some of the changes made, as well as the challenges posed by Open Access publishing and Diversity / Inclusivity concerns. 

There are changes to Council going forward. I’m delighted that Jon Ablett has been nominated as President, with Fiona Allan as Vice President, joining Phil Fenberg in that role. Heartfelt thanks to those who are standing down from Officer positions – Rowan Whittle as meetings Secretary (succeeded by Debbie Wall-Palmer), Jon Ablett as awards Secretary (succeeded by Lauren Sumner-Rooney), Tom White as membership Secretary (succeeded by Harriet Wood). Welcome to Andreia Salvador as Archivist – so no longer a “Void” in that role! There is no named officer for the Forum – should there be? – Phillip Hollyman has done a sterling job of navigating the Forum over the last two years. 

I am grateful to all of Council, as well as those named, for the work you have put into making the _Society_ vibrant and vital. The ‘unique challenges’ of 2020/21 have I think been met, thanks to your efforts and the support given to me – thank you for this. And in closing, a reminder: our most excellent and hard-working Treasurer, Katrin Linse, stands down this year. We need to identify her successor. 

John Grahame, President MLS 

See next page for the Council at March 2021, and nominations going forward to 2022: 

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|**Year of existence**|**2020-2021**|**2021-2022**|
|---|---|---|
||**127**|**128**|
|**President**|John Grahame (3)|Jon Ablett (1)|
|**Vice Presidents**|Robert Cameron(3)|Fiona Allan(1)|
||Phil Fenberg (2)|Phil Fenberg (3)|
|**Ex officio**||John Grahame(1)|
|**Councillors**|Debbie Wall-Palmer(1)|Alan Hodgson(2)|
||Alan Hodgson(1)|Robert Cameron(1)|
||Andreia Salvador(3)|PhillipHollyman(3)|
||Fiona Allan(3)|Aidan Emery (1)|
||PhillipHollyman(2)|Victoria Sleight(1)|
||Lauren Sumner Rooney (2)|Katie Collins(1)|
|**Co-opted**|Aidan Emery|Rowan Whittle(1)|
||||
|**Journal Editor**|Dinazarde Raheem|Dinazarde Raheem|
|**Bulletin Editor**|Georges Dussart|Georges Dussart|
|**Treasurer**|Katrin Linse|Katrin Linse(finalyear)|
|**Membership Secretary**|Harriet Wood|Harriet Wood(1)|
|**Hon. Secretary**|Rowan Whittle|Debbie Wall-Palmer(1)|
|**Web manager**|Tom White/Chong Chen|John Grahame (web)/Chong Chen<br>(Facebook)|
|**Awards Officer**|Jon Ablett|Lauren Sumner Rooney (1)|
|**Archivist**|void|Andreia Salvador(1)|



‘Years’ means ‘years in post’. These are limited as described in the objects of the Society 




The Malacologist 

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## **Molluscs in marine environments** 

**Welcomed by the President of the Society, John Grahame Hosted on Zoom** 


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Registration was free<br>Harriet Woods handled registrations from<br>Cardiff. Harriet.wood@museumwales.gov<br>Registration closed on Friday March 19th<br>**----- End of picture text -----**<br>


organized by the Council, led by John Grahame, supported by Phillip Hollyman, Katrin Linse, Lauren Sumner-Rooney and Harriet Wood'. 



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## **The evolutionary history of molluscs in vent and seep environments** 

## **Crispin T.S. Little** 

School of Earth and Environment, University of Leeds, Leeds, UK E-mail earctsl@leeds.ac.uk 

Molluscs are abundant constituents of modern hydrothermal vents and hydrocarbon seep (aka. methane or cold seeps) sites and this is partially because of the adaptation of many taxa to a symbiotic relationship with chemoautotrophic bacteria in their gills or oesophagus. The taxonomic diversity of molluscs in chemosynthetic environments tends not to be high, but is dominated rather by a few groups, for example bathymodiolin and vesicomyid bivalves, and provannid gastropods. Molecular divergence estimates indicates that some of these diverse taxa are relatively recent arrivals in chemosynthetic environments during the Cenozoic era, and this is supported by their fossil record. However, other groups are older, with Mesozoic origins. Molluscs are present in Palaeozoic vent and seep sites, but these belong to mostly extinct groups, showing that molluscan adaptation to, and extinction within chemosynthetic environments occurred regularly throughout Earth history. 


**The author sampling fossils from an early Cretaceous seep deposit in Svalbard.** 



**Articulated solemyid bivalve in seep carbonate, early Cretaceous, NE Greenland. Handlens = 1.5 cm diame-** 

**The author (seated with yellow helmet right centre) with students in an open mining pit, Cyprus, from where vent** 


## **Saving species from extinction one species at a time** 

## **Darren Fa** 

University of Gibraltar, Europa Point Campus, Gibraltar GX11 1AA Email - darren.fa@unigib.edu.gi 

Darren Fa described the taxonomy, distribution and ontogeny of the endangered limpet _Patella ferruginea_ . He identified the distributions of patellids and mytilids across the straits of Gibraltar and discussed the effects of tidal amplitude on P.ferruginea and its local ecotope, including on artificial habitats such as boulder armour used to protect against shore erosion.  Conservation efforts were discussed in relation to preferred _P.ferruginea_ habitats in the straits. Individual animals were translocated to more benign habitats, with good results for percentage survival compared with resident populations. Fa discussed the possible extension of the species back into the Mediterranean and the importance of stepping stone populations creating genetic connectivity in this process. 







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## **Snails in space: a study of vestibular system plasticity** 

## **Nikolay Aseyev** 

Institute of Higher Nervous Activity and Neurophysiology Russian Academy of Scientists, Moscow, Russia E-mail: aseyev@ihna.ru 

In 2005 NASA and the Russian space agency, Roskosmos, contacted our lab with an invitation to conduct a flight study of gravitational physiology relevance on board an upcoming unmanned satellite Foton-M2. A separate Russian laboratory had to withdraw from participation because of technical reasons. A vice director of the Institute of Biomedical Problems of the Russian Academy of Sciences knew the situation and began to initiate a potential collaboration of NASA scientists and our laboratory .  Time was short, the payload had to be less than 2 kgs in weight, and the project could not interfere with any other projects or the spacecraft. Most importantly, the project had to have clear scientific merit. The head of my lab, Pavel Balaban, had the resources on hand, an energetic and enthusiastic team, and accepted the challenge. Because of the success of the Foton-M2 mission we were invited to participate in the Foton-M3 and Bion-M1 missions. 

We quickly designed the experiments on our model subject, land snail _Helix lucorum_ . These snails are very hardy and resilient, and can survive several weeks without food and water – the conditions expected on the satellite. The weight of the adult snail is about 25 grams, so we were able to launch a reasonably large group of our snail astronauts. Our expectations were correct, and in all three of our orbital missions, we observed no mortality in snails. The goal of the project focused on the adaptation of the vestibular system of snails to near weightlessness conditions on orbit. We used a systems approach to study the vestibular system employing a wide arsenal of methods. In contrast to marine cephalopod and pteropod mollusks, which use the vestibular system to orient themselves in 3D space, land snails use their vestibular system for vertical movements during the day-night cycle and adapting themselves to circadian temperature changes. Also, the land snails use their vestibular sense for detecting dangerous stimuli, and its activation can mean either the snail is in a fall or under attack from a predator. 

In our study to rule out the known confounding factors of an orbital flight and habitat confinement we used 4 groups of additional controls, with different factors controlled, such as temperature, light cycle, food deprivation, congestion and overloading during landing, which was controlled in ground-based control by short centrifugation of snails with 10G. 

Our lab participated in 3 flight missions. The logistics of sample return after landing back to our lab was under development by the two space agencies and out of our control. The Foton-M2 snails arrived at the lab after 40 hrs post-landing on commercial airplane routes. This delay obscured our ability to detect with confidence the snails' response and adaptation to microgravity. Two consequent missions used private aircraft directly from the nearest airport to Moscow, cutting the time to 10-14 hrs post-landing, and thus were more successful. Our story is based on the results of these two missions. Most of the data came from the last mission on the 30-day Bion-M1 satellite, where our space collaborators and we took into account the mistakes of the earlier missions. 

First, we filmed the negative gravitaxis behavior of the ground control and postflight snails. A snail reacts decidedly when its head is directed downward along the vector of gravity such as an opposed head-down tilt, akin to us pitching face forward to the ground, and turns itself in the opposite direction. Then we analyzed the video records and measured the time parameters of the gravitaxis reaction, also the tentacle withdrawal reflex, which was a proxy for a defensive behavior to dangerous vestibular stimulus. In postflight snails, we found significantly shorter latency of the T0 phase of the reaction, simply the time when snails completely retract their tentacles, reflecting an interpreted characteristic of fear they feel. Significant differences between groups were also found in T0 and T4 phases for the length of the tentacles, but not for the angle between tentacles. We made correlation analyses of all measured parameters and found that snails that were slow in the first phases of gravitaxis response were slow in late phases as well. In addition, the correlation of duration of the phases and tentacles length in postflight snails is more correlated with the weight of the animal. The last finding is not surprising, since postflight snails for a month appeared not to feel the weight of their shells. 


To translate the observed postflight behavioral characteristics of the snails with the 

underlying neural commands, we dug deeper into the vestibular sense organs of snails, the statocysts. This is paired organ, like the gravity-sensing otolith organs (utricle and saccule) we have. The equivalent angular acceleration sensors of the semicircular channels we possess are lacking in the snails. The statocyst contains an inertial mass of calcium carbonate grains or statoconia in the form of aragonite surrounded by mechanosensing hair cells and supporting cells. The number of hair cells in this snail species is only 13 cells, which are primary receptor neurons. 

Vestibular adaptation to spaceflight can occur in the peripheral organ itself or in the neural connectivity or both. A popular hypothesis of vestibular changes occurring as the result of exposure to orbital flight, and often discussed in the literature, is an increase in inertial mass in an attempt to restore the "weight" of the statoliths. We used scanning electron microscopy in several postflight snail statocysts to compare with the superficial structure of two control statocysts and found no significant differences at this level of examination. This pointed more to plasticity associated with the sensory neurons 

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## > CONTINUED 

At the 2013 timeframe of the Bion-M1 mission, new high-throughput sequencing techniques became available to researchers, and thus we attempted to study the transcriptomes of statocyst, which is challenging due to the small amount of RNA in this 13-cell organ and lack of genome data for this snail. Finally, we (with our collaborators) had success and found that different peptides are differentially expressed in postflight and control groups. For one of the more diverse neurotransmitter receptors, glutamate, we also found a significant difference between flight and their ground counterparts. 

We then did physiological experiments in snail statocysts. We recorded the static nerve activity during tilts on postflight and control animals. Later we were able to make a detailed analysis of the records. In the static nerve, neural responses of statocyst are visible, and we were able to distinguish responses of one hair cell from the other by action potential or spike parameters. 

In all three missions, the spontaneous or normal background activity of statocysts didn’t change in postflight animals in comparison with control. The total response activity in the static nerve was significantly higher in the postflight group at some time points. In the two successful missions with different duration of exposure to near weightlessness, we conducted comparable protocols and concluded that the time of neural adaptation to microgravity is less than 12 days (the duration of the shorter mission). The postflight snails of Bion-M1 satellite were prepared for recording over a relatively long period of time, about 6 hours. If we split the postflight group into early and late recordings, the critical time point is 20h after landing, when we clearly see the readaptation of statocyst to normal gravity. We then reanalyzed our data from the previous mission, split the postflight snails of the Foton-M3 satellite, and found the same picture there. The earlier we recorded the neural statocyst activity to perturbation stimuli after landing, the greater were the responses. We made the same comparison on the level of single hair cells, sensitive to the preferred direction of the stimuli. We found an even more profound effect of increased responses on postflight statocysts than in total activity. A very similar critical time point of readaptation to normal gravity after landing was found in studies of vertebrate animals and human astronauts, suggesting a common neuronal basis of vestibular system adaptations. 


## **Are some chitons extremophiles, or mere opportunists on the fringe of hostile deepsea habitats?** 

## **Douglas J. Eernisse** 

California State University, Fullerton, USA Email:  deernisse@fullerton.edu 

Eight chiton species were identified in samples from a methane seep off Costa Rica. They are already known from whalefalls, seamounts and other deep sea samples. Douglas J. Eernisse posed several questions (see Fig. 1). Some chiton species were large, reaching several centimetres in length (Fig. 3) . One of the species _, Placiphora velata_ , is an ambush predator and behaves like a Venus Flytrap. Eernisse concluded that seep habitats provided opportunities for  generalist deep sea species, though some of the genera are know to inhabit shallow seas; there is still much to learn about chitons. 



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Figure 2 


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## Number  77 (August 2021) **Solar radiation stress in terrestrial snails — from thermodynamics to microevolution** The Malacologist Page 38 

## **Heinz-R. Köhler, Institute of Evolution and Ecology, University of Tübingen, Germany** 

Heinz-R. Ko hler[1] , Rita Triebskorn[1] , Ulrich Ga rtner[2] , Henri A. Thomassen[1] , Yvan Capowiez[3] , Christophe Mazzia[4] , Thomas Knigge[5] and the 2018 Theba Survey Consortium[6] 

1 Institute of Evolution and Ecology, University of Tu bingen, Tu bingen, Germany 

2 Faculty of Engineering, Esslingen University of Applied Sciences, Esslingen, Germany 

3 INRA, UMR 1114, Avignon, France 

4 IMBNE, UMR 7263, AMU, CNRS, Universite d´Avignon, Avignon, France 

5 UMR-I 02 SEBIO, Universite du Havre, Le Havre, France 

6 Z. Arad, T. Backeljau, J.K.Y. Burmester, Y. Capowiez, L.J. Chueca, M. Coelho da Silva, M.S. Davies, L. Favilli, J. Florit Gomila, J. Heller, T. Knigge, H.-R. Ko hler, S. Krais, A. Lobo da Cunha, G. Manganelli, S. Mazzuca, T. Monsinjon, C. Moreira, G. MorenoRueda, A. Piro, T. Prieto, J. Quintana Cardona, E. Rolan-Alvarez, B. Rowson, M.J. Saffrey, L. Sawallich, M. Schilthuizen, H. Schmieg, A.E. Staikou, H.A. Thomassen, G. Toma s Faci, R. Triebskorn 

## Email: heinz-r.koehler@uni-tuebingen.de 

Land snails, especially in Mediterranean and arid climates, are sometimes exposed to very strong solar radiation. Nevertheless, some species living in these habitats occur in exceptionally high individual densities, suggesting effective adaptations of these animals to heat and drought stress. In addition, these snails may include highly polymorphic species, in which individuals can differ considerably in the pigmentation of their shells. In this context, adaptation mechanisms on the biochemical, physiological, behavioural and morphological levels and their microevolutionary implications can be elucidated by a combination of laboratory, semi-field and field studies. 

Our work concentrates on helicoid species, particularly on the highly polymorphic species _Theba pisana_ . Using highresolution thermography, µMRT, flow modelling, respirometry, biochemistry and field-based microcosms our team addresses the following questions: Under which conditions do land snails predominantly suffer from thermal stress and how can they limit it? Are different pimentation patterns actually biologically relevant in this context? Which conditions overstrain the capacity of biochemical and physiological defence mechanisms? Is there a thermal selection pressure on more intensely pigmented morphs and, if so, under which climatic conditions? And does this have implications for the future distribution of the species and phenotypic variation of _T. pisana_ at the continental scale in the context of climate change? 

In the year 2017, we installed 54 open-top chambers close to Avignon, Southern France, and investigated about 11,000 individuals of _T. pisana_ ( _Ecology & Evolution_ **11** : 1111-1130 [2021]). In 2018, we conducted a Europe- and Middle East-wide survey, which brought together >30 colleagues providing more than 19,000 _T. pisana_ individuals from 172 sites which we used for pigmentation analysis and spatial modelling of morphs. Our studies revealed the particular importance of climatic parameters for the evolution of morphological variation across the Mediterranean and all of Europe, and models allow predictions of the future continent-wide distribution of this species and its morphs as a result of expected environmental warming. 





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## **Quickfire presentations** — 

These were the MSL’s solution to the problem of not being able to have a poster exhibition. Each on-line presentation was for 5 minutes. If no abstract is here, no abstract was made available. Images are screen grabs from the talks, or were supplied by the presenters. 

## **Victor Kang** 

Postdoctoral researcher Evolutionary biomechanics group Department of Bioengineering, Imperial College, London k.kang@imperial.ac.uk> 


**Limpet feeding on algae growing on the shell of its neighbour** 


**A rare sighting of a limpet actively moving. Unlike barnacles and mussels, which remain permanently attached as adults, limpets often move around to feed. This specimen was photographed midtransition from a glass plate to the aquarium** 



**A method was devised in this study to collect limpets with minimal damage to their soft bodies. Here the attachment organ - the pedal sole - is shown on a recently detached specimen, where several unfortunate barnacles were also detached in the process.** 

_**Patella vulgata**_ **adhere with such strength that an individual can easily withstand the weight of a large rock.** 





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## **Obituaries** 

## **Brian Morton 1942 - 2021** 

Brian Morton, malacologist, marine biologist and marine conservationist died on March 28 2021 at Littlehampton, England.28[th] March, 2021. The contribution that Brian made to our knowledge of the anatomy and ecology of bivalves was profound. Over his career he investigated bivalves from a huge spectrum of families, employing meticulous dissection coupled with serial thin sections. Throughout he was notable for use of ink drawings of shells, gross anatomy and microscopic structures seen in thin section with only rare use of photographic images. These studies followed and developed from the tradition of functional morphology established by Sir Maurice Yonge in previous decades. 

His bivalve investigations began with his PhD (1968) study of the biology of the invasive _Dreissena polymorpha_ in a west London reservoir (his PhD supervisor Richard Purchon was, in turn, a former student of Yonge). Interest in dreissenid bivalves later came back into prominence following invasion of _Dreissena_ into the American Great Lakes, the spread of _Mytilopsis_ around the world and the discovery of a cave-living _Congeria_ in Croatia; Brian was involved in all these studies. Similarly, earlier research into _Corbicula fluminea_ in Hong Kong became globally relevant with the spread into numerous freshwater systems. Together with Joseph Britton, Brian published the definitive practical guide to the study of _Corbicula_ . 

Brian was always fascinated by bivalves at the extremes of morphology or ecology. This led to his studies of the anatomy and ecology of coral-boring bivalves in the families Lithophaginae and Gastrochaenidae and also mangrove mud-living bivalves such as _Geloina_ and _Laternula_ .  A long-term interest was in the anatomy and commensal associations of galeommatoidean bivalves with burrowing invertebrates, such as sipunculans, echiuroids and crustaceans. Another focus was the Anomalodesmata, for a while recognised as a distinct sub-class of Bivalvia although molecular results now confirm the group is part of the radiation of heterodont bivalves. They exhibit a range of life 


_**Stirpulina ramosa**_ **(from Morton 2007)** 

styles but the clade of Cuspidariidae, Verticordiidae and Poromyidae lack normal gills and have adopted a predatory, carnivorous diet, feeding suctorialy on small crustaceans. Over the years, Brian described the anatomy and possible food capture mechanisms of a range of these unusual bivalves (most recently with Fabrizio Machado) that live mainly in deeper water. The most bizarre anomalodesmatan bivalves belong in the super- 

_**Galeomma taki**_ **(from Morton, 1973)** 

family Clavagelloidea most notably the tubular ‘watering pot’ shells _Brechites_ , _Nipponoclava_ and _Verpa_ with the flared, perforated anterior ends resembling the rose on garden watering cans while the true shells are embedded into the wall of the tube. Brian investigated their anatomy and hypothesized about the mode of tube formation.  Other clavagelloideans include forms that excavate crypts in limestone and coral and have short funnel shape tubes. Brian rarely found these genera alive but his persistent search of museum collections and rare finds resulted in evolutionary scenarios and a proposed separation into two distinct families Clavagellidae and Penicillidae. These studies culminating in a recent comprehensive review (with F. Machado) of their origin, relationships and evolution. 

Brian had a long-term interest in the eyes of bivalves, beginning with the siphonal eyes of _Laternula_ and continued with descriptions of the structure and distribution of various types of pallial and cephalic eyes in a wide range of species. Major review papers discussed their evolution and phylogenetic distribution. 

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The ecological importance of Mytilidae in rocky shore communities of Hong Kong led Brian with many students to long term studies of all aspects of the biology of intertidal mussels particularly _Perna_ , _Mytilisepta_ , _Septifer_ and _Brachidontes_ .. Most recently, collaborative molecular studies of Mytilidae have clarified disputed relationships of some species. Following his BSc and PhD at Chelsea College, University of London, Brian worked briefly on shipworms at the Hayling Island Laboratory of University of Portsmouth before moving to the Zoology Department at Hong Kong University in 1971 where he stayed until retirement. It is fair to say that in 1971 HKU was a backwater of marine biology but by the time of Brian’s retirement and led by his efforts it had been transformed into a powerful and influential centre in Asia. Along with the many students that were trained and went on to prominent positions in marine biology or environmental agencies in Hong Kong or abroad, his most notable achievement was the construction and opening in 1990 of the Swire Institute of Marine Sciences at Cape d’Aguilar where he was Director for 13 years. The laboratory has acted as a major focus for marine research ever since. Soon after arriving in Hong Kong Brian realised that on these diverse shores there was a severe taxonom- **Brian collecting 1986 Marine Biological Workshop, Hong Kong** ic impediment that hampered research. In order to address this problem, he organised a series of resi- 

dential malacological and marine biological workshops and invited experts in various animal groups from around the world to visit Hong Kong and conduct field-based research at a youth centre near Tolo Channel in the New Territories. The workshops increased in size and became logistically complex with organised shore work, vehicles, boats, dive groups, trawling and dredging coupled with improvised laboratories. These workshops were amazingly effective with the results published in a series of voluminous proceedings from 1979 –1989. Some of the locations, habitats and their biota documented by the early workshops 

have now disappeared or impacted by increasing urbanisation and the publications stand as a valuable data archive.  Many participants returned regularly to Hong Kong to continue research. Later, three smaller-scale workshops were based at the Cape d’Aguilar laboratory. 

A milestone in marine ecology of Hong Kong was the book ‘The Seashore Ecology of Hong Kong’ (1983) written by Brian and co-author John Morton (from New Zealand and no relation). All the major habitats were described and the constituent flora and fauna illustrated largely by composite line drawings.  Better than 


dry academic research papers this book proved to be major stimulus to education and public awareness of the marine habitats of Hong Kong. Other books on seashore ecology written and illustrated in similar style followed featuring the Azores and Gulf of Mexico.  A lifelong connection to the Azores with repeated research visits began in 1965 with an undergraduate expedition from Chelsea College. There Brian first met and became friends with Antonio Frias Martins (then a trainee priest) who later became a zoologist and they collaborated on the book ‘Coastal ecology of the Azores’ (1998).  Another book ‘Shore ecology of the Gulf of Mexico’ (1989) resulted from a long-term research collaboration and friendship with Joseph Britton then of Texas Christian University. In the 1970s the marine environment of Hong Kong was severely impacted by pollution and habitat destruction from increasing urbanisation. Soon after arriving Brian became a passionate, vociferous and outspoken advocate for the formation of marine reserves and controls of the marine disposal of industrial and domestic effluent. Change came slowly but in the end persistence paid off with the establishment of a series of marine parks and reserves including the corals at the Hoi Ha Marine Park and the Cape d’Aguilar reserve.  The first and major success was the World Heritage site of the Mai Po wetlands administered by the World Wildlife Fund. 



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By any standards his research output in terms of publications was enormous and sustained by prodigous drive and determination throughout all his career. In retirement in 2003 and back at Littlehampton the stream of papers, books and articles continued and he was still completing papers in the last weeks of his life. For all the many achievements his constant need to keep proving himself or outdoing perceived competitors either within his own institute or more broadly was at times challenging for friends and colleagues.   Although often scathing in criticism of the work of others he could be belligerent when his own work was challenged. Some of us remember Friday evening consumption of alarming quantities of beer in the Senior Common Room at Hong Kong University which brought out both the best and worst in a generous, talented and complex man. 

Brian received many awards for his conservation work including election to the Global 500 by UNEP, investiture as Knight of the Golden Ark, Netherland (1997), Order of the British Empire (1999) and the Duke of Edinburgh Conservation Gold Medal (2004). He was a long-term supporter of the Malacological Society and elected as Honorary Life Member. 

John Taylor   (with input from Emily Glover and Liz Harper) 


## **John B. (“Jack”) Burch 1929 - 2021** 

I am deeply saddened to inform you that our esteemed colleague John B. (“Jack”) Burch died yesterday at the age of 92 after a long illness. He had been very well taken care of by his family and passed away peacefully in their company in Littleton, Colorado. His devoted partner of 69 years, Peggy Burch (pegburch@umich.edu), informed me that a family memorial or Celebration of Life may be held at a later date (we are still in the middle of a pandemic) and that, in lieu of flowers, donations in his memory be made to The John B. Burch Malacology Fund, or to a charity of your choice. Jack had an extraordinary, long-lived, and highly influential career in science serving as Professor and Curator of Molluscs at the University of Michigan since 1962 ( _Emeritus_ since 2001). See below an outline of some of the highlights, and his _Curriculum Vitae_ for details. 

## **Honors & Awards** 

Honorary Life Memberships: American Malacological Society (2009), Malacological Society of Korea (1994), Malacological Society of the Philippines, (1994). Lifetime Achievement Award: The Freshwater Mollusc Conservation Society (1999). Special Recognition Award: The Science Society of Thailand (1995). 

## **Research Interests** 

Jack had broad interests in the biology of non-marine Mollusca, especially freshwater snails, and he made numerous significant contributions to their global study. Early in his career, a major focus was the application of karyo- 


**In English pub 2015 Published by Mackie** 


logical, serological, and tissue culture techniques to characterize the roles of intermediate snail hosts in the epidemiology of human schistosome parasites. This involved extensive field work throughout the tropics and lab work in Ann Arbor (below, left). One of his major immuno-cytological discoveries was the presence of cryptic diploid and polyploid (tetra/ hexa/octoploid) lineages in populations of the important intermediate host _Bulinus truncates/tropicus_ complex, that were differentially susceptible to infection by human schistosome parasites. This seminal work led to years of research (supported by NIH, NSF, and the World Health Organization) and international outreach, the latter involving numerous workshops on medical malacology as well as the training of scientists from a large diversity of countries. 

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In addition to his medical malacology research, Jack made many important basic science contributions to our fundamental knowledge of non-marine molluscan diversity in his >180 publications. He was also distinguished by his prodigious outreach and service to malacology as a discipline. One aspect of this was his exquisite guides to North American freshwater molluscs ( _Freshwater Unionacean Clams of North America, Freshwater Sphaeriacean Clams of North America, North American freshwater snails_ ) and land snails ( _Land Snails of The University of Michigan Biological Station Area, Identification of Eastern North American Land Snails, How to Know Eastern Land Snails_ ). Another was his astonishing record of founding four new malacological journals, one of which, _Malacologia_ (currently the top-ranked Zoological journal by citation metrics), endures. The other three are now either transformed into a new entity ( _Walkerana_ , now _Freshwater Mollusc Biology and Conservation_ ), or, after decades of production ( _Malacological Review_ , _Journal of Medical and Applied Malacology_ ), retired. 

To paraphrase Shakespeare ( _Othello_ ), Jack has “ _done the state_ (of Malacology) _some service, and they know’t. “_ He will be missed. 


**Jack Burch in the 1960s** 

Diarmaid O  Foighil 

Museum of Zoology & Department of Ecology & Evolutionary Biology The University of Michigan. 

Editor’s note -In an email conversation with Diarmaid about Jack Burch, I told him that in my younger days, I’d met Jack  at conferences a few times and found him to be a gentlemanly, kind person. Diarmaid replied “ Your interaction with Jack was highly characteristic. He was a warm and supportive personality who wore his scientific pre-eminence lightly and was invariably encouraging to younger scientists - that was also my experience. “ 


## **Arie W. Janssen** 

With great sadness we inform you that Arie W. Janssen, nestor of fossil and recent malacological research in the Netherlands, passed away in the night of 6th August 2021. He reached the age of 84 years and during the last few months suffered from a rapidly deteriorating physical condition; nonetheless until just a few days before his death he kept on working on pending publications with zeal and a bright mind as ever. 

Arie was a self-taught researcher and had a long career as malacologist and palaeontologist that started at the Rotterdam Museum of Natural History in the mid nineteen sixties. From 1969 onwards he continued at the Rijksmuseum van Geologie en Mineralogie (the National Museum of Geology and Mineralogy) that is nowadays part of Naturalis Biodiversity Center in Leiden. As curator of the fossil mollusc collections he ensured major enlargements through fieldwork and through donations from the networks Arie had been building. The collections at the department of fossil mollusca rank among the top ones of their kind in the world. Arie’s extensive field work and research during the first part of his career was focused on Tertiary and Quaternary molluscs mainly from the North Sea Basin, but also from other parts of Europe. 

Since the mid nineteen eighties,  Arie increasingly concentrated his research on fossil planktic gastropods (mostly pteropods and heteropods). He promoted and implemented the stratigraphic use of these groups of fossil molluscs, as they proved to be very useful to enable 


**The front of Arie's funeral card, depicting a pteropod of the genus** _**Edithinella.**_ **Arie described and named it after his wife Edith.It also shows his favourite definition of pteropoda, which he used in most of his messages and emails combined with his signature.** 

long-distance correlations and the establishment of stratigraphic ages of geological units. He developed into a specialist of worldwide renown in this field and personally contributed to what is today the world’s largest planktic molluscan collection (fossil + recent), housed in Naturalis. Nowadays planktic gastropods are important indicators for the state of our oceans and their increasing acidity. Arie’s research is being continued by the Naturalis Plankton Group under the leadership of Katja Peijnenburg. 

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Arie was one of the co-founders of the WTKG, the Dutch Work Group of Tertiary and Quaternary Geology, a team of professional and above all non-professional enthusiasts. He built a widespread network that delivered important results in scientific ‘grassroots’ research of fossil molluscs, but also on local stratigraphy. Besides encouraging the members with his thorough knowledge and ability to enthuse people, Arie was also for many years the editor of the WTKG’s scientific publications, that appeared from 1964 first under the name _Mededelingen_ (Contributions) and from 2002 onwards under the name _Cainozoic Research_ . 

Arie leaves behind an enormous œuvre of more than 300 publications, smaller papers but also a considerable number of voluminous books. A highlight for the knowledge of Dutch Miocene molluscs was his 450-pages book with a 82-plates atlas on the fossil fauna of the Miste beds (near Winterswijk, The Netherlands) that appeared in 1984. But not only fossil molluscs caught his interest; in 1965 he wrote, together with the recently deceased Ed de Vogel, a book on Dutch freshwater molluscs. This early work found its updated continuation in the 1998 standard work _De Nederlandse Zoetwatermolluscen_ , authored by a consortium of scientists with Arie’s participation. But his most important scientific legacy are the many papers with groundbreaking research results on planktic molluscs. After his retirement in 1997, Arie and his wife Edith moved to Malta, where he intensified his research on fossil pteropods. They moved back to The Netherlands in 2013, where Arie continued his research work even up to the last days of his admirable and fruitful life. His name will live on through no less than 32 eponyms, named in honour of his work by colleague scientists. Besides twenty-three new mollusc and five new fish species, four new mollusc genera bear his name. But above all his name Janssen will remain forever present, connected to no less than 175 molluscan species described by him as new to science, a stunning 115 of which in the Pteropoda and Heteropoda, plus one new family, one new subfamily, twelve new genera and two new subgenera. 

He will be greatly missed by colleagues and worldwide co-reseachers. It is a great loss for his friends and to science, and above all to his wife, his son and his daughter and his four grandchildren. 

Jaap van der Voort (close friend and independent reseacher, Ostercappeln, Germany), with contributions from 

Frank Wesselingh, Ronald Pouwer and Katja Peijnenburg (Naturalis Biodiversity Center, Leiden, The Netherlands) 

LIST OF TAXA DESCRIBED BY ARIE W. JANSSEN (INCLUDING with CO-AUTHORS) SORTED BY YEAR OF PUBLICATION, AND SHOWING THE TAXONOMY USED  IN THE ORIGINAL PUBLICATION, IRRESPECTIVE OF POSSIBLE LATER EMENDATIONS.             Newly described (sub)genera or (sub)families are shown in _CAPITALS_ . 

_Solariella (Solariella) formosa_ Janssen, 1967 _Skenea minuta_ Janssen, 1967 _Daronia (?Cyclostremella) punctata_ Janssen, 1967 _Circulus quadricarinatus_ Janssen, 1967 _Cingula (Ceratia) regiorivi_ Janssen, 1967 _Cingula (Chevallieria) pseudoproxima_ Janssen, 1967 _PSEUDALVANIA gen.nov._ Janssen, 1967 _Pseudalvania dingdensis_ Janssen, 1967 _Teinostoma (Solariorbis) hosiusi_ Janssen, 1967 _Teinostoma (?Solariorbis) partimstriatum_ Janssen, 1967 _Gegania miocenica_ Janssen, 1967 _Cerithiopsis (Cerithiopsis) vogeli_ Janssen, 1967 _Cerithiopsis (Cerithiopsis) vandermarki_ Janssen, 1967 _Cerithiopsis (Cerithiopsis) andersoni_ Janssen, 1967 _Cirsotrema (Opaliopsis) turbonillaeforme_ Janssen, 1967 _Cirsotrema (?Opaliopsis) koeneni_ Janssen, 1967 _Circulus praecedens gliberti_ Janssen, 1967 _Putilla gottscheana westfalica_ Janssen, 1967 _Aclis (Stilbe) neglecta_ Janssen, 1969 _Polinices (Euspira) staringi_ Janssen, 1969 _Phalium (Semicassis) bicoronatum belgicum_ Janssen, 1969 _ldasola lignicola_ Janssen, 1969 _Axinulus germanicus_ Janssen, 1969 _Alvania (Actonia) basisulcata_ Janssen, 1969 _Amaea (Undiscala) marialuisae_ Janssen, 1969 _Turriscala (s.l.) straeleni germanica_ Janssen, 1969 _Euspira edithae_ Janssen, 1969 _Hinia (Tritonella) twistringensis_ Janssen, 1969 _Cancellaria (Merica) contorta gelriana_ Janssen, 1969 _Narona (Aneurystoma) canaliculata_ Janssen, 1969 

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## >CONTINUED 

_Microdrillia teretiaeformis_ Janssen, 1969 _Aphanitoma ronaldi_ Janssen, 1969 _Aphanitoma fransi_ Janssen, 1969 _Conus (Chelyconus) altenai_ Janssen, 1969 _Eulimella (Eulimella) lutaria_ Janssen, 1969 _Ebala (Ebala) vandervlerki_ Janssen, 1969 _Spiratella kautskyi_ Janssen, 1969 _Astarte (Isocrassina) omalii scaldensis_ Janssen, 1974 _Tridonta (Tridonta) domburgensis_ Janssen, 1974 _Gyraulus (Gyraulus) acuticarinatus vleminckxae_ Janssen, 1980 _Nystia (Nystia) glibertheinzeli_ Janssen, 1980 _Pododesmus (Monia) anitae_ Janse & Janssen, 1983 _Skenea schuermanni_ Janse & Janssen, 1983 _Rissoella (Jeffreysiana) hesselinki_ Janse & Janssen, 1983 _Hinia (Hinia) cimbrica_ (Ravn, 1907) _voorthuyseni_ Janse & Janssen, 1983 _Babylonella stemerdinki_ Janse & Janssen, 1983 _Daphnella defectiva_ Janse & Janssen, 1983 _MISTEIA_ gen.nov. Janssen, 1984 _Trigonostoma (Misteia) mistense_ Janssen, 1984 _Trigonostoma barnardi_ Janssen, 1984 _Trigonostoma lindeni_ Janssen, 1984 _Trigonostoma pouwi_ Janssen, 1984 _Trigonostoma geslini josephinae_ Janssen, 1984 _Sveltia gliberti_ Janssen, 1984 _GLIBERTTURRICULA_ gen.nov.  Cade e & Janssen, 1985 _Glibertturricula vervoeneni_ Cade e & Janssen, 1985 _Spirulirostra baetensi_ Janssen & Mu ller, 1985 _CAPEDOPECTEN_ gen.nov. Dijkstra & Janssen, 1988 _Clio gailae_ Goedert & Janssen, 2020 _VAGINELLINAE_ subfam.nov. Janssen 2020 _Capedopecten anellus_ Dijkstra & Janssen, 1988 _Limacina ingridae_ Janssen, 1989 _Limacina irisae_ Janssen, 1989 _Limacina jessyae_ Janssen, 1989 _Limacina mariae_ Janssen, 1989 _Linacina wilhelminae_ Janssen, 1989 _‘Creseis’ berthae_ Janssen, 1989 _Clio blinkae_ Janssen, 1989 _Clio irenae_ Janssen, 1989 _Clio jacobae_ Janssen, 1989 _Clio pauli_ Janssen, 1989 _SPOELIA_ gen.nov. Janssen, 1990 _Spoelia torquayensis_ Janssen, 1990 _Limacina curryi_ Janssen, 1990 _Limacina lunata_ Janssen, 1990 _Limacina tatei_ Janssen, 1990 _Vaginella victoriae_ Janssen, 1990 _Vaginella sannicola_ Janssen, 1990 _Clio nielseni_ Janssen, 1990 _Vaginella tricuspidata_ Zorn & Janssen. 1993 _FREDENIA_ gen.nov. Cade e & Janssen, 1994 _Streptodytion impiger_ Cade e & Janssen, 1994 _Streptodyction schnetleri_ Cade e & Janssen, 1994 _. Streptodyction twistringensis_ Cade e & Janssen, 1994 _Streptolathyrus masculinus_ Cade e & Janssen, 1994 _Streptolathyrus regularis_ Cade e & Janssen, 1994 _Fusus dhondtae_ (nom.nov.) Cade e & Janssen, 1994 _Gamopleura melitensis_ Rehfeld & Janssen, 1995 _Cuvierina jagti_ Janssen, 1995 _Ireneia marqueti_ Janssen, 1995 _Ireneia nieulandei_ Janssen, 1995 _Clio giulioi_ Janssen, 1995 _DIACROLINIA_ gen.nov. Janssen, 1995 _? Diacrolinia elioi Janssen, 1995 EDITHlNELLA_ gen.nov. Janssen, 1995 _SPHAEROCINIDAE_ fam.nov. Janssen & Maxwell, 1995 

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## CONTINUED > 

_Philine aquila_ van der Linden & Janssen, 1996 _Edithinella curva_ Janssen, 1998 _Diacrolinia larandaensis_ Janssen, 1999 _STRIOLIMACINA_ nom.nov. Janssen, 1999 _Clio lucai_ Janssen, 2000 _Clio (Balantium) collina_ Janssen & Zorn, 2001 _Brocchinia gerdae_ Janssen & Petit, 2003 _CURRYLIMACINA_ (gen.nov.) Janssen, 2003 _Heliconoides vanderweideni_ Janssen, 2004 _Bowdenatheca miocenica_ Janssen, 2004 _Clio (s.lat.) ghawdexensis_ Janssen, 2004 _Edithinella bonaviai_ Janssen, 2004 _BELLARDICLIO_ subg.nov. Janssen, 2004 _Cavolinia landaui_ Janssen, 2004 _Cavolinia vendryesiana_ f. _Pliomediterranea_ Janssen, 2004 _Cuvierina (Cuvierina) pacifica_ Janssen, 2005 _URCEOLARIA_ subg.nov. Janssen, 2005 _Cuvierina (Urceolaria) cancapae_ Janssen, 2005 _Cuvierina (Urceolaria) curryi_ Janssen, 2005 _Ireneia gracilis_ Janssen, 2005 _JOHNJAGTIA_ gen.nov. Janssen, 2005 _Vaginella basitrunctata_ Janssen, 2005 _Edithinella doliarius_ Janssen, 2006 _Urceolarica_ nom.nov. Janssen, 2006 (for _Urceolaria_ Janssen, 2005) _Atlanta lingayanensis_ Janssen, 2007, _Atlanta richteri_ Janssen, 2007, _Atlanta seapyi_ Janssen, 2007, _Heliconoides sondaari_ Janssen, 2007, _Striolimacina andaensis_ Janssen, 2007, _Hyalocylis marginata_ Janssen, 2007, _Cavolinia baniensis_ Janssen, 2007, _Cavolinia perparvula_ Janssen, 2007, _Cavolinia shibatai_ Janssen, 2007, _Diacavolinia pristina_ Janssen, 2007, _Diacria italica_ Grecchi, 1982 f. _fissicostata_ Janssen, 2007 _Diacria microstriata_ Janssen, 2007, _Diacria paeninsula_ Janssen, 2007, _Diacria philippinensis_ Janssen, 2007 _Sphaerocina convolvula_ Janssen, 2007 _Heliconoides lillebaeltensis_ Janssen, Schnetler & Heilmann-Clausen, 2007 _Heliconoides linneensis_ Janssen, 2008 _HAMECONIA_ gen.nov. Janssen, 2008 _Hameconia edmundi_ Janssen, 2008 _Heliconoides daguini_ Cahuzac & Janssen, 2010, _Heliconoides merlei_ Cahuzac & Janssen, 2010, _Heliconoides pyrenaica_ Cahuzac & Janssen, 2010 _Limacina_ ? _vegrandis_ Cahuzac & Janssen, 2010 _Creseis antoni_ Cahuzac & Janssen, 2010 _Vaginella gaasensis_ Cahuzac & Janssen, 2010 _Clio lozoueti_ Cahuzac & Janssen, 2010 _Clio vasconiensis_ Cahuzac & Janssen, 2010 _Diacrolinia cluzaudi_ Cahuzac & Janssen, 2010, _Creseis roesti_ Cahuzac & Janssen, 2010 _Heliconoides mermuysi_ Cahuzac & Janssen, 2010 _Creseis tugurii_ Cahuzac & Janssen, 2010 _Peracle charlotteae_ Janssen & Little, 2010 _Limacina erasmiana_ Janssen, 2010 _Limacina guersi_ Janssen, 2010 _Gamopleura maxwelli_ Grebneff, Janssen & Lee, 2011 _Limacina asiatica_ Janssen, King & Steurbaut, 2011 _Limacina dzheroiensis_ Janssen, King & Steurbaut, 2011 _Clio (Clio) pyramidata_ Linne, 1767 f. _tyrrhenica_ A.W. Janssen, 2012 _Protatlanta kbiraensis_ Janssen, 2012 _Carinaria maempeli_ Janssen, 2012 _Heliconoides wardijaensis_ Janssen, 2012 _Limacina ernstkittli_ Janssen, 2012 _Creseis curta_ Janssen, 2012 

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## >CONTINUED 

_Styliola schembriorum_ Janssen, 2012 _Johnjagtia baharensis_ Janssen, 2012 _Clio merijni_ Janssen, 2012 _Clio vilis_ Janssen, 2012 _Cavolinia gatti_ Janssen, 2012 _Cavolinia microbesitas_ Janssen, 2012 _Diacrolinia pumilionis_ Janssen, 2012 _Gamopleura pilula_ Janssen, 2012 _Peracle amberae_ Janssen, 2012 _Peracle grebneffi_ Janssen, 2012 _Clione? ignotus_ Janssen, 2012 _Clione? imdinaensis_ Janssen, 2012 _Clione? phosphoritus_ Janssen, 2012 _Clione? tripartitus_ Janssen, 2012 _Clione? tumidulus_ Janssen, 2012 _Heliconoides vonhachti_ Janssen, 2012 _Tibiella watupuruensis_ Janssen, Renema & Wesselingh, 2013 _Altaspiratella tavianii_ Janssen, Jagt, Yazdi, Bahrami & Sadri, 2013 _Limacina aryanaensis_ Janssen, Jagt, Yazdi, Bahrami & Sadri, 2013 _Limacina perforata_ Janssen, Jagt, Yazdi, Bahrami & Sadri, 2013 _Limacina yazdii_ Janssen, Jagt, Yazdi, Bahrami & Sadri, 2013 _Texacuvierina hodgkinsoni_ Janssen, Jagt, Yazdi, Bahrami & Sadri, 2013 _ZHGENTIANA_ gen.nov. A.W.Janssen, R.Janssen & van der Voort, 2015 _Cuvierina tsudai_ Burridge, Janssen & Peijnenburg, 2016 _Limacina novacaesarea_ Janssen & Sessa, 2016 _Limacina tanzaniaensis_ Janssen, 2017 _Heliconoides nikkieae_ Janssen, 2017 _Limacina timi_ Janssen, 2017 _Limacina parvabrazensis_ Garvie & Janssen, 2020 _Limacina pseudopygmaea_ Garvie & Janssen, 2020 


______________________________________________________________________________________________ 

## **Adhesive defence mucus secretions in the red triangle slug (** _**Triboniophorus graeffei**_ **) can incapacitate adult frogs** 

Gastropods secrete mucus for a variety of purposes, including locomotion, reproduction, adhesion to surfaces, and lubrication. A less commonly known function of mucus secretion in this group involves its use as a defence against predation. Among the terrestrial slugs, mucus that serves this particular purpose has been studied for only a handful of species under laboratory conditions, where it is thought to be produced for self-fouling or to make individuals difficult to consume. However, the mechanisms of how these defensive secretions operate and their effectiveness in deterring predation in the natural world have not be described in much detail. Adhesive mucus secretions in the red triangle slug ( _Triboniophorus graeffei_ ) may be an adaptation against predation. Field observations of a large redeyed green tree frog ( _Litoria chloris_ ) trapped in the mucus secretions of a nearby _T. graeffei_ revealed that this mucus serves to incapacitate predators rather than just simply as an overall deterrence. Mechanical stimulation of _T. graeffei_ under laboratory conditions revealed that adhesive secretions were produced from discrete sections of the dorsal surface when disturbed, leading to the production of a highly sticky and elastic mucus that was unlike the thin and slippery mucus used during locomotion. The adhesiveness of the defensive secretions was strengthened and reactivated when in contact with water. This appears to not only be the first description of defensive mucus production in this slug species but one of the first natural observations of the use of adhesive defence secretions to incapacitate a predator. The biomechanical properties of this mucus and its ability to maintain and strengthen its hold under wet conditions make it potentially useful in the development of new adhesive materials. 

Gould, J., Valdez, J.W. & Upton, R. email: jose.valdez@bios.au.dk 

bioRxiv preprint doi: https://doi.org/10.1101/544775 




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## **Forthcoming meetings** 

## **The Malacological Society of London** 

http://www.Malacsoc.org.uk 

## **The Molluscan Forum** 

Thursday 18t[h] November 2021 0945h  – 1630h (to be confirmed) 

**CALL FOR REGISTRATIONS AND PAPERS DEADLINE 16 OCTOBER 2021** 

This informal, annual, and successful meeting is designed to bring together people starting their research on molluscs, to give them the opportunity to present and discuss their work and to compare notes on methods and problems. The deadline for registrations and talk applications is 


Attendance to the Molluscan Forum is open to all, but preference is given to **research students, postpost-doctoral researchers, undergraduate students starting molluscan projects, and amateurs** engaged in substantial projects that have not yet been published. Any topic related to molluscs is acceptable: palaeontological, physiological, behavioural, ecological, systematic, morphological, cellular, or molecular. 

Short talks (~15 mins) or quick fire talks (3 slides, 5 mins, in lieu of posters) may be offered. They need not be polished accounts of completed work; descriptions of new methods, work in progress, and appeals for assistance with unsolved problems are equally acceptable. 

## **There is NO registration fee.** 

## **Enquiries and registrations to:** 

Phil Hollyman, Fisheries Ecologist, British Antarctic Survey (phyman@bas.ac.uk) 

**Non-presenters:** please let us know you will be coming so that we can estimate numbers. 

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>CONTINUED 

# **The Malacological Society of London** 

# **Molluscan Forum, Thursday 18t[h] November 2021 9:45 am – 4.30 pm** 

## **REGISTRATION FORM** 

## **Return before 1[st] October 2021, by email to:** 

## Phil Hollyman, Fisheries Ecologist, British Antarctic Survey (phyman@bas.ac.uk ) 

Name................................................................................................................…………… 

Address..........................................................................................................…………..… 

......................................................................................................................……………… 

Tel. No................................................................................................…………................. 

Email.................................................................................................……………............... 

**Status:** Research Student / Undergraduate / Post-doctoral researcher / amateur (delete as appropriate) 

‘Other’ (please state) ……………………….. 

Short talks (~15 min) or posters may be offered. 

Instead of posters, we will have 5 minute (3 slide maximum) quick-fire powerpoint presentations, 

I wish to give a talk / quick-fire talk (delete as appropriate) entitled: 

.............................................................................................................................……………………… 

..............................................................................................................................……………………... 

**Please attach, as a Microsoft Word attachment, an abstract of not more than 350 words, TOGETHER WITH TWO .JPG IMAGES IN SUPPORT OF THE ABSTRACT.   Abstracts and images of accepted contributions will be published in the Society’s on-line bulletin which is called** _**The Malacologist. The Malacologist**_ **has an ISSN number and is published and archived on the website of the MSL.** 

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>CONTINUED 

## **Abstract submission** 

Abstracts submitted for the Molluscan Forum should be sent as Microsoft Word files. 

Please use the following format: 

Title (12pt, left justified) <blank line> 

Authors (10 pt, left justified, presenting author underlined; use superscript numbers to indicate institutional affiliation) 

<blank line> 

Institutions (10pt, left justified; in this order: Number (superscript), Department, Institution, City, Country) Presenting Author email <blank line> Abstract (11pt, no indentation, left justified, 350 words maximum) 

## **EXAMPLE ABSTRACT** 

## **The geographic scale of speciation in** _**Stramonita**_ **(Neogastropoda: Muricidae)** 

## **Martine Claremont[1,2] , Suzanne T. Williams[1] , Timothy G. Barraclough[2] , and David G. Reid[1]** 

1Deptartment of Zoology, Natural History Museum, London, UK 2Department of Biology, Imperial College London, Berkshire, UK Email: m.claremont@nhm.ac.uk 

_Stramonita_ is a relatively small, well-defined genus of muricid marine gastropods limited to the tropical Eastern Pacific and the Atlantic. The type species, _S. haemastoma_ , is known to have teleplanic larvae and is estimated to remain in the water column for several weeks. _Stramonita haemastoma_ shows regional variation, and this has led to the recognition of five geographical subspecies: _S. h. haemastoma_ , from the Mediterranean and Eastern Atlantic to Brazil, _S. h. floridiana_ , on the east coast of Florida and in the Eastern Caribbean, _S. h. caniculata_ on the west coast of Florida and the Gulf of Mexico, _S. h. rustica_ in the Western Caribbean and _S. h. biserialis_ in the Eastern Pacific. The protoconch has been shown to be similar across the _S. haemastoma_ complex, implying that all subspecies have equally long lived larvae. Within these subspecies, cryptic variation is suspected. For example, _S. h. biserialis_ is suggested to be differentiated North/ South on a small scale. In the presence of teleplanic larvae, speciation on such a small scale seems paradoxical. Various explanations for this paradox are possible. Actual (or realized) dispersal of _Stramonita_ species may be more limited than presently believed, leading to allopatric differentiation. Alternatively, morphological differentiation may not be a reliable indicator of genetic differentiation, and _S. haemastoma_ ( _sensu lato_ ) might indeed prove to be a single taxon. It is also possible that ecological speciation could result in geographical speciation on a small scale in the presence of wide dispersal. My results suggest that five species of _Stramonita_ are present in the Caribbean, at least three of which occur sympatrically. Gene flow is maintained between Caribbean and Mediterranean populations in at least one species, while no genetic differentiation was found along the Eastern Pacific coast. The implications of these results are discussed. 

**NOTE THAT ABSTRACTS ARE PUBLISHED IN** _**THE MALACOLOGIST**_ **WHICH  IS THE BULLETIN OF THE SOCIETY AND HAS AN ISSN NUMBER.** 

**BEFORE THE FORUM, PLEASE EMAIL TO THE EDITOR OF** _**THE MALACOLOGIST**_ **(Email address below) TWO IMAGES TO ACCOMPANY YOUR ABSTRACT. TRY TO MAKE THESE IMAGES ONES THAT YOU WOULD NOT USE IN AN EVENTUAL FULL PAPER.** 

**EDITOR        georges.dussart@canterbury.ac.uk** 




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## **Grants and Awards** 

## **Malacological Society of London Awards and Grants** 

The Research Awards Scheme was established to commemorate the Society's Centenary in 1993. Under this scheme, the Society gives awards to support research on molluscs that is likely to lead to publication.  The closing date for applications each year is 15th December.  Grants are preferentially conferred on students and researchers without regard to nationality or membership of the Society. Preference is also given to discrete research projects that fall within the subject areas covered by the Society's _Journal of Molluscan Studies_ . Applications will be assessed by scientific merit, value of the project and for student applicants, the extent to which the research will benefit the applicant's scientific aspirations. The successful applicants will be notified by 31st March and announced at the Annual General Meeting.  Awardees are encouraged to publish their work in the _Journal of Molluscan Studies_ (full papers) or _The Malacologist_ (travel award reports, research award reports, news of ongoing research etc) as appropriate, 

## _**Early Career Research grants**_ 

Eligibility is restricted to those investigators at the outset of their independent scientific career. Applications must therefore be 1) postgraduate students, 2) within five years of being awarded their PhD (adjustable for career breaks), or 3) independent researchers not having a PhD. Early Career Research Grants will only be awarded to individuals twice, but not within 3 years of receiving a first award 

## _**Sir Charles Maurice Yonge Award**_ 

There is no application process for Sir Charles Maurice Yonge Awards. These awards are given for the best Travel Award application on bivalves, by a member of the Society to attend an international meeting (not including the Molluscan Forum). Authors of exceptional studies on bivalves in _the Journal of Molluscan Studies_ may on occasion also be given this award. The Editor will nominate such papers as he/she sees fit. The award covers the costs requested in a Travel Award, or for open access publication of the paper.  Members of the Society will also receive a personal cash prize of £300. Non-members will receive a personal cash prize of £250 plus one year’s membership to the Society. If a paper is multi-authored, the award will be made to the corresponding author. 

## _**Senior Research Awards**_ 

are aimed at established researchers in professional positions, but without regard to nationality. Applicants for Senior Research Awards must be members of the Malacological Society of London. The Society currently awards up to five Senior Research Grants per year, each with a value of up to £1,500, to support research on molluscs that is likely to lead to publication. The maximum amount available should not be considered as a ‘target’; rather requests should reflect the research that is proposed. The grants are reviewed by a Reviewers Panel including both Council and non-Council members invited for that purpose. 

## _**Travel Grants**_ 

Travel Awards are available as bursaries to support attendance at a conference or workshop relevant to malacology. Grants are preferentially conferred on students but researchers without professional positions may also apply. The maximum amount for one of these awards is £500 for Society members and £300 for non-members.  Preference will be given to members of the Society. There are two closing dates each year, 30th June for travel starting between 1st September of the current year and 28th February of the following year, and 15th December for travel starting between 1st March and 31st August of the following year. 

For further information, guidance notes and to access the application form see here - http://malacsoc.org.uk/awards-andgrants/travel-grants 

## _**Annual Award**_ 

This Award is made each year for an exceptionally promising initial contribution to the study of molluscs. This is often a thesis or collection of publications. The value of the Award is £500. Candidates need not be a member of the Society but must be nominated by a member. There is no application form: the nominating member should send the material for evaluation with a covering letter or letter of support to the Honorary Awards Secretary. The closing date each year is 1st November. The winner(s) will be notified by 31st March, and announced at the Annual General Meeting. 

## _**Applications**_ 

Applications for Research Awards and Travel Grants should be sent to the Honorary Awards Secretary, Jonathan Ablett, Division of Invertebrates, Department of Life Sciences, Natural History Museum, London, SW7 5BD  For further information, guidance notes and to access the grant application form see  http://malacsoc.org.uk/awards-and-grants/research-grants Please note that all applications must be sent by email to MSL_awards@nhm.ac.uk. 




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## **Malacological Society of London—Membership notices** 

## **Objects** 

The objects of the Society are to advance education and research for the public benefit by the study of molluscs from both pure and applied aspects. We welcome as members all who are interested in the scientific study of molluscs. There are Ordinary Members, Student Members and Honorary Members. Members are entitled to receive a digital copy of the _Journal of Molluscan Studies_ and such circulars as may be issued during their membership. The society's Web Site is at: 

http://www.Malacsoc.org.uk 

## **Publications** 

The Society has a continuous record of publishing important scientific papers on molluscs in the _Proceedings_ , which evolved with Volume 42 into the _Journal of Molluscan Studies_ . The _Journal_ is published in annual volumes consisting of four parts which are available on -line by members and student members.  The Society no longer produces paper copies of the Journal. Members also receive  on-line access to _The Malacologist_ , which is the bulletin of the Society,  issued twice a year, in February and August. _The Malacologist_ is published  on the website of the Society. 

## **Meetings** 

In addition to traditional research on molluscan biology, physiological, chemical, molecular techniques are amongst the topics considered for discussion meetings and papers for publication in future volumes of the _Journal_ . 

## **Subscriptions** 

## **Membership fee structure** 

Ordinary Members: Journal  on-line only    £45 Student Members:   Journal  on-line only    £25 

## **Methods of Payment** 

- (1) Sterling cheque  to "The Malacological Society of London". 

- (2) Banker’s standing order to: HSBC (Sort code 40-16-08 Account no. 54268210)   63-64 St Andrew’s Street, Cambridge C32 3BZ 

- (3)  Overseas members  wishing to pay electronically should should use 

IBAN   GB54MIDL4016084268210 

SWIFT/BIC  MIDL GB22 

(4) Credit card: Overseas members ONLY may pay by credit card: the Society can accept VISA and MasterCard payments only. Please provide the Membership Secretary with your card number and expiry date, card type (VISA or MasterCard.), the name on the card, and the cardholder’s address (if this differs from your institutional address). Receipts will only be sent if specifically requested. 

## **Institutional Subscriptions to the Journal** 

Enquiries should be addressed directly to Oxford University Press, Walton Street, Oxford OX2 6DP, U.K. 

## **Change of Member's Address** 

## **Please inform the Membership Secretary of a change of postal or email address** 


## ——————————————————————————————————————————————— 

## **APPLICATION FOR MEMBERSHIP OF THE MALCOLOGICAL SOCIETY OF LONDON** 

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**REGISTERED CHARITY NUMBER: 275980** 

**REPORT OF THE TRUSTEES AND UNAUDITED FINANCIAL STATEMENTS FOR THE YEAR ENDED 31 DECEMBER 2020 FOR THE MALACOLOGICAL SOCIETY OF LONDON** 

Staffords Chartered Accountants Unit 1, Cambridge House Camboro Business Park Oakington Road, Girton CAMBRIDGE Cambridgeshire CB3 0QH 



**THE MALACOLOGICAL SOCIETY OF LONDON** 

**CONTENTS OF THE FINANCIAL STATEMENTS FOR THE YEAR ENDED 31 DECEMBER 2020** 

||Page|
|---|---|
|Report of the Trustees|1 - 3|
|Independent Examiner's Report|4|
|Statement of Financial Activities|5|
|Balance Sheet|6|
|Notes to the Financial Statements|7 to 11|





**THE MALACOLOGICAL SOCIETY OF LONDON** 

**REPORT OF THE TRUSTEES FOR THE YEAR ENDED 31 DECEMBER 2020** 

The trustees present their report with the financial statements of the charity for the year ended 31 December 2020. The trustees have adopted the provisions of the Charities SORP (FRS102) ‘Accounting and Reporting by Charities: Statement of Recommended Practice applicable to charities preparing their accounts in accordance with the Financial Reporting Standards applicable in the UK and Republic of Ireland (FRS 102)(effective 1 January 2015)’, Financial Reporting Standard 102 ‘The Financial Reporting Standard applicable in the UK and Republic of Ireland’ and the Charities Act 2011. 

## **STRUCTURE, GOVERNANCE AND MANAGEMENT** 

## **Governing document** 

The charity is controlled by its governing document, a deed of trust, and constitutes an unincorporated charity. 

The governing document is continually reviewed following its original adoption on 17 May 1978, with the current operative revision of the governing document being that amended on 28 April 2005. 

## **Recruitment and appointment of new trustees** 

Nominations for Council (the board of related party) from members must be proposed and seconded and in the hands of the Secretary by December 31st. Members at the AGM 2016 proposed and seconded that only the Society’s President and Officers (Treasurer, Secretary, Membership Secretary, Awards Secretary) are forming the board of trustees.  It shall be the duty of Council to nominate members for election to the offices of President, Treasurer, Secretary, Membership Secretary, Editor of the Journal, Editor of the Bulletin, Archivist, Web Manager, and Awards Secretary, and for the vacancies in the Council caused by annual retirement. Nominations from the members and from Council shall be submitted to the Society with the notice convening the Annual General Meeting which shall be sent to every member of the Society not less than fourteen days before the Meeting. 

In the case of a vacancy arising in any office of the Society, or in the Council, other than by way of resignation or retirement in the Annual General Meeting, the Council shall have power to appoint a temporary Council member to that vacancy until the next Annual General Meeting. 

## **Risk management** 

The trustees have a duty to identify and review the risks to which the charity is exposed and to ensure appropriate controls are in place to provide reasonable assurance against fraud and error. 

## In particular: 

(a) Admission fees and annual subscriptions shall be such sums as may be determined by a Special General Meeting convened under Rule VI(c), or at the Annual General Meeting. 

(b) Subscriptions shall be due on the 1st of January in each year; but in the case of a new member, immediately after election. Members elected during the months of November and December, however, shall not be required to pay for the year in which they are elected. 

(c) Any member whose current subscription has not been received in full by 31 January shall be reminded of the arrears in subscription and be informed that despatch of the Journal is suspended until the arrears are paid. No member whose subscription is twelve months in arrears shall be entitled to any of the privileges of the Society, and any member whose subscription is two years in arrears shall cease to be a member of the Society, unless the Council decide otherwise. 

(d) The Council shall revise and publicise the conditions of the Society's awards and grants from time to time. 

(e) For the purpose of legal protection of the property of the Society, all funds, books and other property shall be declared vested in Council as the Society's Trustees. 

(f) The Council shall cause to be kept Minutes of Council and Society Meetings and books of account in respect of all receipts, payments, assets and liabilities. Accounts shall be presented to each Annual General Meeting for approval by members and such accounts shall be audited or independently examined as appropriate. 

Page 1 



**THE MALACOLOGICAL SOCIETY OF LONDON** 

**REPORT OF THE TRUSTEES FOR THE YEAR ENDED 31 DECEMBER 2020** 

## **OBJECTIVES AND ACTIVITIES** 

## **Objectives and aims** 

The governing document contains the following in relation to the objective of the charity: 

The objects of the Society are to advance education, research and learning for the public benefit in the study of molluscs from both pure and applied aspects. In furtherance of these objects, but not further or otherwise, the Society shall have the following powers: 

(a) To promote and co-ordinate meetings and symposia, 

(b) To promote and co-ordinate research both pure and applied; 

(c) To provide for the worldwide dissemination of the useful results of such research by publication of the Journal of Molluscan Studies; 

(d) To award prizes to outstanding students in the field of molluscan biology; 

(e) To award research grants to individuals which will advance the study of molluscan biology; 

(f) To do all such things as will further the objects of the Society. 

## **Significant activities** 

In 2020 the Malacological Society of London had proposed meeting at the Marine Biological Association in Plymouth on "Limpets 2020" that had to be cancelled with a week’s notice due to Covid-19, and the annual Molluscan Forum for Young Scientists was held virtually. The Society published the members' bulletin "The Malacologist" and in cooperation with Oxford University Press "Journal of Molluscan Studies". 

## **Public benefit** 

The charity's objects are for the public benefit because increasing public knowledge is required as they form an important part of the global biodiversity and ecosystem stability which can have effects on human health and are a human food source. 

## **ACHIEVEMENT AND PERFORMANCE** 

## **Charitable activities** 

In 2020 the main charitable activities were the funding of eleven research projects, of which nine claimed the grants (total cost of £12,262), eight to young researchers and malacologists in non-permanent positions and one to senior researchers, the support of 3 students and young researchers to attend conferences and training courses, an activity highly impacted by Covid-19 as the Society usually supports around 20 scientists here. 

## **FINANCIAL REVIEW** 

The finances of the Malacological Society have been pleasing during 2020 with an overall gain of £52,220. This gain is explained by a gain in the Fixed Interest and Investment funds and lower awards and meeting expenditure. 

Our investments had an overall gain of £28,997 (comparing market value at 31 December 2020 with market value at 31 December 2019), with the COIF Investment Fund making a gain of £25,132 and the COIF Fixed Interest Fund a gain of £3,865. During 2020, no funds were transferred from the current account to savings accounts. 

Separately, the profit-share from the publication of the Journal of Molluscan Studies in 2020 provided the Society with most of its income contributing £48,890. The Editor of the Journal, Dr Dinarzarde Raheem, and the Assistant Editors are to be commended for their hard work contributing to the publication of our scientific journal. In addition, sales of the digital archives provided £507 of income. 

In 2020, more funds were used for research awards, being £12,262 in 2020 compared to £8,696 in 2019, while travel awards remained similar, but there was reduced spending on Council meeting and Forum travel awards as meetings after January 2020 were held virtually. The Society (MSL) spent less money in 2020 compared to 2019, this was mainly based on less expenses paid for meetings and to the JMS editor for attendance expenses, Malacological conferences to promote our journal and network with potential assistant editors and authors. 

Page 2 



**THE MALACOLOGICAL SOCIETY OF LONDON** 

**REPORT OF THE TRUSTEES FOR THE YEAR ENDED 31 DECEMBER 2020** 

## **REFERENCE AND ADMINISTRATIVE DETAILS Registered Charity number** 275980 

**Principal address** c/o British Antarctic Survey High Cross Madingley Road CAMBRIDGE CB3 0ET 

**Trustees** Dr J Grahame President Dr T White Membership Secretary, elected 4[th] June 2020 Dr K Linse Hon. Treasurer Dr R Whittle Hon. Secretary J Ablett Awards Officer Dr T White Ex-Membership Secretary, term ended 4[th] June 2020 

**Independent examiner** Steven Ellis FCA Staffords Chartered Accountants Unit 1, Cambridge House Camboro Business Park CAMBRIDGE Cambridgeshire CB3 0QH Approved by order of the board of trustees on ............................................. and signed on its behalf by: 

............................................. Dr John Grahame - Trustee 

Page 3 



**INDEPENDENT EXAMINER'S REPORT TO THE TRUSTEES OF THE MALACOLOGICAL SOCIETY OF LONDON** 

I report on the accounts for the year ended 31 December 2020 set out on pages five to ten. 

## **Respective responsibilities of trustees and examiner** 

The charity's trustees are responsible for the preparation of the accounts. The charity's trustees consider that an audit is not required for this year (under Section 144(2) of the Charities Act 2011 (the 2011 Act)) and that an independent examination is required. 

It is my responsibility to: 

- examine the accounts under Section 145 of the 2011 Act 

- to follow the procedures laid down in the General Directions given by the Charity Commission (under Section 145(5)(b) of the 2011 Act); and 

- to state whether particular matters have come to my attention. 

## **Basis of the independent examiner's report** 

My examination was carried out in accordance with the General Directions given by the Charity Commission. An examination includes a review of the accounting records kept by the charity and a comparison of the accounts presented with those records. It also includes consideration of any unusual items or disclosures in the accounts, and seeking explanations from you as trustees concerning any such matters. The procedures undertaken do not provide all the evidence that would be required in an audit, and consequently no opinion is given as to whether the accounts present a 'true and fair view ' and the report is limited to those matters set out in the statements below. 

## **Independent examiner's statement** 

In connection with my examination, no matter has come to my attention: 

- (1) which gives me reasonable cause to believe that, in any material respect, the requirements 

   - to keep accounting records in accordance with Section 130 of the 2011 Act; and 

   - to prepare accounts which accord with the accounting records and to comply with   the accounting requirements of the 2011 Act 

have not been met; or 

- (2) to which, in my opinion, attention should be drawn in order to enable a proper understanding of the accounts to be reached. 

Steven Ellis FCA Staffords Chartered Accountants Unit 1, Cambridge House Camboro Business Park CAMBRIDGE Cambridgeshire CB3 0QH 

Date: ............................................. 

Page 4 



## **The Malacological Society of London** 

**Statement of Financial acitivities  (incorporating an income and expenditure account) for the year ended 31st December 2020** 

|Note<br>**INCOMING RESOURCES**<br>**Income from Activities of the Charity**<br>Members’ Subscriptions: Current Year<br>Collection of Subscription Arrears<br>4<br>OUP: Income from Journal Publication<br>2<br>Income from sale of Digital Archive<br>Sundry Income<br>Donations<br>**Income from Investments**<br>3<br>Interest<br>CCLA<br>3<br>Dividends<br>**TOTAL INCOMING RESOURCES**<br>**RESOURCES USED**<br> **Awards**<br>Annual Award<br>WCM Award<br>7<br>Research Awards (Early career & Senior)<br>EuroMal 2020 award<br>7<br>Travel Bursaries for conferences<br>5,6<br>Travel awards: AGM<br>5,6<br>Travel awards: Forum<br>**Directly Relating to Work of Charity**<br>4<br>Journal expenses<br>4<br>Journal colour plates<br>4<br>Journal editor expenses, incl meetings<br>5<br>Bulletin Expenses<br>6<br>Meeting Expenses<br>Independent examiners expenses<br>5,6<br>Council Meeting travel expenses<br>Web sites<br>5<br>Postage, Printing & Stationary<br>Charges for cc subscription collection<br>Bank charges<br>Sundries<br>**Total**<br>**NET INCOME**<br>8<br>GAIN ON REVALUATION OF FIXED ASSETS<br>**NET MOVEMENT IN FUNDS**<br>TOTAL FUNDS BROUGHT FORWARD<br>**TOTAL FUNDS CARRIED FORWARD**|TOTAL<br>2020<br>1,670<br>**‐**<br>48,890<br>507<br>66<br>**‐**<br>6<br>3,706<br>**54,844**<br>**‐**<br>**‐**<br>12,262<br>**‐**<br>1,050<br>1,240<br> **‐**<br>**14,552**<br>4,637<br>8,280<br>1,644<br>41<br>358<br>1,128<br>738<br>**‐**<br>**‐**<br>18<br>222<br>4<br>**31,621**<br>**23,223**<br>28,997<br>52,220<br>391,090<br>**443,310**|TOTAL<br>2019|
|---|---|---|
|||2,335<br>‐<br>48,620<br>1,351<br>100<br>20<br>12<br>3,706|
|||**56,143**|
|||‐<br>100<br>8,696<br>1,800<br>2,100<br>412<br>2,070|
|||**15,178**<br>4,750<br>5,220<br>5,285<br>148<br>5,663<br>1,080<br>1,795<br>‐<br>‐<br>37<br>204<br>2|
|||**39,363**<br>**16,780**<br>49,217<br>65,997<br>325,093<br>**391,090**|



## **CONTINUING OPERATIONS** 

All income and expenditure has arisen from continuing activities. 

Page 5 



## **The Malacological Society of London** 

## **Balance Sheet at 31 December 2020** 

|Note||||2020|||2019|
|---|---|---|---|---|---|---|---|
|||||£|||£|
||**FIXED ASSETS**|||**400,619**|||**371,622**|
|8|Tangible  Assets|||||||
||**CURRENT ASSESTS**|||||||
|11|Debtor|||‐|||‐|
|9|Cash at Banks|||55,819|||32,548|
|10|CREDITORS: (Amounts falling due within one year)||‐|13,128<br>||‐|13,080<br>|
||**NET CURRENT ASSETS**|||**42,691**|||**19,468**|
||**TOTAL NET ASSETS**|||**443,310**|||**391,090**|
||**FUNDS**|||||||
||Unrestricted (Designated):|||||||
|11|Annual Award Fund|||6,500|||6,500|
|11|CM Yonge Award Fund|||9,250|||9,250|
|11|Centenary Research Award Fund|||80,000|||80,000|
|||||95,750|||95,750|
|11|General reserve Fund|||347,560|||295,340|
||**TOTAL FUNDS**|||**443,310**|||**391,090**|
||Katrin Linse|John Grahame||||||
||Honorary Treasurer  & Trustee|President  & Trustee||||||
||Dated|Dated||||||



Page 6 



## **The Malacological Society of London** 

## **Notes to the Financial Statements for the year ended 31st December 2020** 

## **1. ACCOUNTING POLICIES** 

## **Basis of Preparing the Financial Statements** 

The financial statements have been prepared in accordance with the Charities SORP (FRS 102) 'Accounting and Reporting by Charities: Statement of  Recommended Practice applicable to charities preparing their accounts in accordance with the Financial Reporting Standards applicable in the UK and Republic of Ireland (FRS 102)(effective 1 January 2015)', Financial Reporting Standard 102 'The Financial Reporting Standard applicable in the UK and Republic of Ireland' and the Charities Act 2011. 

‐ The SORP normally requires a columnar fomat, in this entities case the trustees' view is this is not appropriate as there are neither restricted funds nor endowment funds and so all of the activity in the year would fall within the unrestricted fund column. 

## **Accounting Convention** 

The financial statements have been preprared under the historical cost convention, except for those items described immediately below. 

‐ The historic cost basis of accounting is used except for investments, which have been included at their market value where readily available at the yearend date. 

## **Going concern** 

The Trustees consider that there are no material uncertainties about the charity's ability to continue as a going concern. 

## **Reconciliation with previously Generally Accepted Accounting Practice** 

In preparing the accounts, the trustees have considered whether in applying the accounting policies required by FRS102 and the Charities SORP FRS 102 the reinstatement of comparative items was required.  In the case of this charity, no changes were made. 

## **Fund accounting** 

‐ The Society funds, including funds available for awrds, are not subject to any restrictions regarding their use, and are available to be used for the general purposes of the Charity. Consequently they are classified as 'Designated Funds'. 

## **Taxation** 

‐ The charity is exempt from tax on its charitable activities. 

## **Significant judgements** 

Apart from those judgements involving estimations, the management has not made any judgements in the process of applying the entity's accounting policies that have significant effect on the amounts recognised in the accounts.  There are no key assumptions concerning the future or other key sources or estimation uncertainty at the reporting date that have significant risk of causing a material adjustment to the carrying amounts of assets and liabilities within the next reporting period. 

## **Financial reporting standard 102 ‐ reduced disclosure exemption** 

The charity has taken advantage of the following disclosure exemption in preparing these financial statements, as permitted by FRS 102 'The Financial Reporting Standard applicable in the UK and Republic of Ireland': 

## **‐** 

the requirements of Section 7 Statement of Cash Flows 

## **Resources Expended** 

Expenditure is accounted for on an accruals basis and has been classified under headings that aggregate all cost related to the category.  Where costs cannot be directly attributed to particular headings they have been allocated to activities on a basis consistent with the use of resources. 

Page 7 



## **The Malacological Society of London** 

## **Notes to the Financial Statements for the year ended 31st December 2020 (cont'd)** 

## **Incoming Resources** 

All income is recognised in the Statement of Financial Activities once the charity has entitlement to the funds, it is probable that the income will be received and the amount can be  measured reliably. 

## **Foreign currencies** 

‐ Asessts and liabilities in foreign currencies are translated into sterling at the rates of exchange ruling at the balance sheet date. Transactions in foreign currencies are tranlated into sterling at the rates of exchange ruling at the date of transaction. Exchange differences are taken into account in arriving at the operating result. 

## **Incoming Resources and Resources used** 

∙ Subscriptions for annual membership are treated as follows: 

|Pre‐paid in prior year|Liabilities|
|---|---|
|Paid in current year|Incoming Resource: subscriptions|
|Unpaid at 31st December|Not included in accounts. No debtor|
|Subsequently recieved|Incoming Resource: collection of subscriptions|
||in arrears|



∙ The costs of the Society Journal and supplements are written off in the year of initial distribution. 

∙ The Profit Share from Oxford Journals is included in the year of receipt and not accrued. 

The amount of Profit Share cannot be determined accurately by OUP until after the Society's AGM. 

## **2. SALE OF THE DIGITAL ARCHIVE BY OUP** 

Since 2010 the sale of the digital archives by OUP has generated a windfall profit of £13,969 in 2010, £9,284 in 2011, £7,299 in 2012,  £10,068 in 2013,  £7827 in 2014, £10,590 in 2015, £2,805 in 2016, £10,429 in 2017,  £4238 in 2018, and £1352 in 2019. 

Although the main customers for this product have now purchased the archive and OUP has split archive sales into subjects, digital archive sales are still ongoing  with £507 in 2020. 

## **3. INTERESTS & DIVIDENDS** 

Income has been derived from the following sources and has been applied to funds as indicated: 

|Sources<br>COIF Deposit Fund Interest<br>COIF Fixed Interest Fund Dividend<br>COIF Investment Fund Dividend<br>Beneficiary<br>Revenue Fund<br>Annual Award Fund<br>CM Yonge Award Fund<br>Research Grants other Funds|2020<br>2019<br>£<br>£<br>6<br>12<br>3,706<br>3,706|
|---|---|
||**3,712**<br>**3,718**|
||1,179<br>1,181<br>194<br>194<br>273<br>273<br>2,066<br>2,070|
||**3,712**<br>**3,718**|



## **4. JOURNAL** The surplus resulting from publication of the Journal is as follows 

|Profit Share from OUP<br>Less: sales of digital archives<br>Profit Share from OUP re Journal<br>Less:printing costs<br>plates<br>Editor & CM meeting expenses<br>Under/over‐provision in previous year<br>**Surplus on publication of the Journal**|2020<br>2019<br>£<br>£<br>49,397<br>49,971<br>507<br>‐<br>1,352<br>‐|
|---|---|
||**48,890**<br>**48,619**<br>12,000<br>‐<br>12,000<br>‐<br>8,280<br>‐<br>5,220<br>‐<br>1,644<br>‐<br>5,285<br>‐<br>6,858<br>7,250|
||**33,824**<br>**33,364**|



Page 8 



**The Malacological Society of London** 

## **Notes to the Financial Statements for the year ended 31st December 2020 (cont'd)** 

## **5. RELATED PARTY TRANSACTIONS AND TRUSTEES' EXPENSES** 

No Trustee or Council Member has recieved any renumeration during the current or previous year.  Expenses paid by Trustees and Council Members on behalf of the Society have been reimbursed during the current year as follows. 

|Trustee<br>Date<br>£<br>Raheem<br>21.01.20<br>20.00<br>Raheem<br>21.01.20<br>812.00<br>Wood<br>21.01.20<br>85.50<br>Fenberg<br>22.01.20<br>23.95<br>Cameron<br>22.01.20<br>45.80<br>Hodgson<br>22.01.20<br>50.80<br>Grahame<br>22.01.20<br>67.90<br>Grahame<br>22.01.20<br>237.89<br>Linse<br>22.01.20<br>23.45<br>Hollyman<br>22.01.20<br>26.90<br>Whittle<br>22.01.20<br>23.45<br>Dussart<br>22.01.20<br>26.45<br>Wall‐Palmer<br>22.01.20<br>107.00<br>Sumner‐Rooney<br>22.01.20<br>18.60<br>Raheem<br>15.06.20<br>812.00<br>Grahame<br>15.06.20<br>14.39<br>Hollyman<br>09.11.20<br>148.79<br>Grahame<br>14.12.20<br>14.39<br>Dussart<br>15.12.20<br>40.68<br>**XPENSES**<br>costs have been incurred on meetings for the Society:<br>AGM:<br>Speakers travel<br>General<br>Dinner<br>Forum:<br>Travel<br>Reception drinks<br>Zoom<br>General 2018 (invoiced in 2019)<br>Benugo (General 2019)<br>Council:<br>Travel<br>Zoom|Reason<br>WCM travel<br>JMS edits<br>Council meeting travel<br>Council meeting travel<br>Council meeting travel<br>Council meeting travel<br>Council meeting travel<br>Council meeting travel<br>Council meeting travel<br>Council meeting travel<br>Council meeting travel<br>Council meeting travel<br>Council meeting travel<br>Council meeting travel<br>JMS edit support<br>Zoom licence CM<br>Zoom licence Forum<br>Zoom licence CM<br>Malacologist<br>2020<br>2019<br>£<br>£<br>1,240<br>412<br>‐<br>183<br>‐<br>204<br>‐<br>2,070<br>‐<br>158<br>149<br>‐<br>‐<br>2,419<br>‐<br>2,700<br>738<br>1,795<br>15<br>‐|
|---|---|
||**2,126**<br>**9,940**|



## **6. MEETING EXPENSES** 

The following costs have been incurred on meetings for the Society: 

Page 9 



## **The Malacological Society of London** 

## **Notes to the Financial Statements for the year ended 31st December 2020 (cont'd)** 

## **7. GRANTS AND AWARDS TO INSTITUTIONS** 

Although grants and awards are given to individuals, in many cases those individuals are affiliated with an institution. Under the SORP, the Society is required to give an analysis of the range of institutions for whom grants and awards are paid: 

£ 

Early Career Res. Grants: 1,420 University of Aberdeen (UK) 1,500 Assoziazone Cheph (It) 1,100 Keio University (Jp) 1,460 University of Bilbao (Spain) 1,402 University of Exceter (UK) 1,457 University of Adelaide 1,500 University of St Andrews (UK) 1,500 University of Turin (It) 

Senior Research Grants 923 Clarkson University  (US) 

In the year the total amount of grants to institutions was £12,262.25. 

## **8. INVESTMENTS** 

In  2020 no funds were transferred to the COIF Investment fund and  to the Fixed Interest Fund. The following investments are held as at 31st December 2020 

||Number|Price|Market|Book|
|---|---|---|---|---|
||of units||Value|Value|
|||31‐Dec‐20|31‐Dec‐20|31‐Dec‐19|
|||£|£|£|
|COIF Investment Fund|1,364.59|206.603|281,928|256,796|
|(Accumulation Units)|||||
|COIF Fixed Interest Fund|83,479.39|1.4218|118,691|114,826|
||||**400,619**|**371,622**|



These investments have been valued at market price (£400,619) in the Balance sheet, with appropriate adjustment (£28,997  ‐ £0 transfers) for the increase in their value (£28,997) in the Statement of Financial activities as an unrealised gain. 

## **9. CASH DEPOSITS** 

The following accounts are held and the balances in each account are: 

|Northern Bank Current Account<br>COIF Deposit<br>Pay Pal<br>HSBC Bank Current Account<br>Total|2020<br>2019<br>£<br>£<br>‐<br>‐<br>2,101<br>96<br>‐<br>‐<br> <br>53,718<br>30,452|
|---|---|
||**55,819**<br>**30,548**|



Page 10 



## **The Malacological Society of London** 

## **Notes to the Financial Statements for the year ended 31st December 2020 (cont'd)** 

## **10. CURRENT ASSESTS & LIABILITIES** 

The following debtors are outstanding 

|||||2020|2019|||
|---|---|---|---|---|---|---|---|
|||||£|£|||
|Forum Travel awards not used||||‐<br>‐||||
|The following creditors are outstanding:||||||||
|||||2020|2019|||
|||||£|£|||
|Society Journal (provision)||||12,000|12,000|||
|Accruals||||1,128|1,080|||
|||||**13,128**|**13,080**|||
|**11. UNRESTRICTED FUNDS**||||||||
|The following movements have taken place within the||Society's four||designated funds:||||
||Revenue|Annual||CM<br>Reseach &|||Total|
|||Award||Yonge<br>other||||
|||||Award<br>Awards||||
||£||£|£|£||£|
|**Balance: 1st January 2020**|**295,340**||**6,500**|**9,250**|**80,000**||**391,090**|
|Interest Earned|1,178||194|273<br>|2,066||**3,711**|
|Grants|‐|‐||‐<br>‐|15,632<br>|**‐**|**15,632**<br>|
|Allocated Surplus: 2020|51,042|‐|194<br>|273<br>‐|13,566||**64,141**|
|**Balance: 31st December 2020**|**347,560**||**6,500**|**9,250**|**80,000**||**443,310**|



The Trustees can, by resolution at one of their meetings, re‐designate these funds for other purposes. 

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