Addition 2 to Géominpal Belgica. 5.2.: Presence of tubes of teredinid

Transcription

Addition 2 to Géominpal Belgica. 5.2.:
Presence of tubes of teredinid-like bivalves (Mollusca)
in some petrified pieces of wood
discovered in the Sint-Niklaas Phosphorite Bed
(Marine Belgian Oligocene),
with some reflections concerning
the Systematic of the Orders Myoida and the Anomalodesmata.
By
Jacques Herman1, Hilde Van Waes1,
Julien Van Nuffel2, Jacqueline Cloetens2,
Eric Vanderhoeft4 and Thierry Vanderhoeft5
1
Herman J. & Van Waes H. : Mail: jacquesalbertherman@gmail.com
Van Nuffel J. & Cloetens J. : Rue Jules La Haie 203, 1090 (Jette, Belgium)
4
Vanderhoeft E. : Rue Simonis 31, 1050 (Ixelles, Belgium)
5
Vanderhoeft T. : t.vander@skynet.be
2
S.V.K. Clay Pit 4: B.G.S. Archives: N°: 42 W 394:
Tubes of Teredinid Bivalvia (Mollusca) in a piece of petrified wood.
Origin: Belsele (Western Flanders, Belgium)
Scheerders Van Kerchove Claypit N°4
Sint Niklaas Phosphorite Bed.
(Lower marine Oligocene)
Diameter of the wood piece: 84 millimetres.
Photographer: Hugues Doutrelepont.
HERMAN Jacques Editor
H
Dedication:
This work is dedicated to the memory of
Hilde, Louise, Rachel Van Waes
05.10.1950 (Sint-Niklaas) – 25.06.2015 (Brussels)
Philologist in Germanic languages - V.U.B. (1972),
honorary citizen (Museology) of Sint-Niklaas-Waes,
Professor of English and Dutch to the Belgian School of Kinshasa
and subsequently to the Flemish Atheneum of Hoboken, Etterbeek and Vilvoorde,
co-editor of
Elasmobranches and Stratigraphy, Taphonomy of some Belgian Cenozoic levels,
co-author of Géominpal Belgica
and
the wife who contemplated with the senior-author
more than thirteen thousand
sunrises and sunsets.
In her garden at Beigem, in August 1995.
Her husband Jacques Herman,
and her friends:
Eric Vanderhoeft, Thierry Vanderhoeft,
Julien Van Nuffel and Jacqueline Cloetens.
At Beigem, 25 Juny 2015
Jacques Herman,Eric Vanderhoeft, Thierry Vanderhoeft,
Julien Van Nuffel and Jacqueline Cloetens.
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Table of Contents
1. Summary - Résumé - Samenvatting - Kurzfassung - Resumen - Resumo - Резюме: p.: 5.
2. Introduction: p.: 7.
3. Introduction: p.: 7.
4. Introductie: p.: 7.
5. Phylum Mollusca: p.: 8.
5.1. Systematics: p.: 8.
5.2. Remarks: p.: 8.
6. Class Bivalvia: p.: 8.
6.1. Generalities: p.: 8.
7. Family Teredinidae: p.: 9.
7. 2. Sub-Family Teredinae: p.: 9.
7. 3. Sub-Family Kuphinae: p.: 15.
7. 4. Sub-Family Bankiinae: p.: 15.
7.5. Geographical distribution of the Family Teredinidae: p.: 19.
7.6. Geological range of the Family Teredinidae: p.: 19.
7.7. Ichnology and the Teredinidae: p.: 19.
7.8. References for Family Teredinidae: p.: 20.
8. Family Xylophagaidae: p.: 24.
8.2. Geographical distribution of the Family Xylophagaidae: p.: 25.
8.3. Geological range of the Family Xylophagaidae: p.: 25.
8.4. Ichnology and the Xylophagaidae: p.: 25.
8.5. References for Family Xylophagaidae: p.: 25.
9. Data furnished by the pieces of wood: p.: 26.
9.1. Nature of these pieces of wood: p.: 27.
9.2. Degradation of these pieces of wood: p.: 27.
9.3. Duration of their drifting: p.: 27.
9.4. First embedding: p.: 27.
9.5. Power of the dis-embedding flow: p.: 27.
9.6. Filling of the teredinid tubes: p.: 27.
9.7. Second embedding: p.: 28.
9.8. Geochemical alteration of this filling: p.: 28.
10. Conclusions: p.: 28.
10.1. Systematic conclusion: p.: 28.
10.2. Taphonomical interpretation: p.: 29.
11. Special adaptations of some Bivalvia: p.: 29.
11.1. General considerations: p.: 29.
11.2. Identic or different valves: p.: 29.
11.3. Advantages presented by the possession of identic valves: p.: 29.
11.4. Advantages presented by the possession of different valves: p.: 30.
11.5. Disadvantages presented by the by the possession of different valves: p.: 30.
12. Short examination of the principal data concerning the diverse Families
phylogenetically considered as parents of the Teredinidae: p.: 30.
12.1. Motivation: p.: 30.
12.2. Recent revision : p.: 30.
12.3. Remarks: p.: 31.
12.4. Family Myidae: p.: 31.
12.5. Family Myochamidae: p.: 32.
12.6. Family Corbulidae: p.: 32.
12.7. Family Pandoridae: p.: 33.
12.8. Family Lanternulidae: p.: 34.
12.9. Family Lyonsiidae: p.: 35.
12.10. Family Periplomatidae: p.: 36.
12.11. Family Thraciidae: p.: 37.
12.12. Family Cuspidariidae: p.: 37.
12.13. Family Verticordiidae: p.: 39.
12.14. Family Teredinidae and Family Xylophagaidae: p.: 40.
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12.15. Family Hiatellidae: p.: 40.
12.16. Family Pholadidae: p.: 44.
12.17. Family Gastrochaenidae: p.: 45.
12.18. Family Clavagellidae: p.: 46.
12.19. Family Penicillidae: p.: 47.
12.20. Family Pholadomyidae: p.: 47.
13. Conclusions: p.: 48.
13.1. Generalities: p.: 48.
13.2. Concerning the systematic position of the extant and extinct teredinid taxa: p.: 48.
13.3. Concerning the systematic position of the fossils
discovered in the Sint-Niklaas Phosphorite Bed: p.: 48.
13.4. Concerning the present conception of the generic taxa
of the extant Myoidea and Anomalodesmata: p.: 49.
13.5. Concerning the present conception of the Order Anomalodesmata: p.: 49.
14. Plates 1 to 42: p.: 53.
15. Comments on the Plates: p.: 95.
16. Acknowledgements: p.: 101.
Remind
Electronic Series do not need Index
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Summary
(Hilde Van Waes)
On examining for the first time, the fossils stored in the drawer: SVK - Vegetal Remains, it was surprising to
realize that some of the silicified pieces of wood contained tubes of Teredinidae (Bivalvia, Mollusca), other ones
traces of activities of larvae of xylophagous insects (Coleoptera, Insecta) and that the eleven elongated siderite
concretions could be fragments of tubes of sabellid-like animals (Sabellida, Polychaeta, Annelida).
This Géominpal Belgica is the second addition to the Publication* which focused on the study of the remains of
Invertebrata discovered in the Sint-Niklaas Phosphorite Bed of Rupelian Age (Belgian Lower Oligocene). This
fascicule is devoted to the examination of the teredinid tubes present in the majority of these petrified pieces of
wood.
*HERMAN, J. & VAN WAES, H. 2013: Géominpal Belgica. 5(2).
Keywords: Belgium, Oligocene, Rupelian, Sands of Ruisbroek, Sint-Niklaas Phosphorite Bed, Teredinidae,
Xylophagaidae, Terediniformes, Myoidea, Anomalodesmata, Bivalvia, Mollusca, Invertebrata.
Résumé
(Jacques Herman)
En examinant, pour la première fois, les fossiles rangés dans le tiroir: SVK - Restes végétaux, grande fut la
surprise de constater que certains morceaux de bois silicifiés étaient porteurs de tubes de Teredinidae (Bivalvia,
Mollusca), d’autres de traces d’activités de larves d’insectes xylophages (Coleoptera, Insecta) et que les onze
concrétions sidéritiques, y rangées, pouvaient être des fragments de tubes d’animaux sabelliformes (Sabellida,
Polychaeta, Annelida).
Ce Géominpal Belgica constitue la seconde addition à la Publication* ayant eu pour thème l’étude des restes
d’Invertebrata découverts dans l’Horizon à Phosphorites de Sint-Niklaas d’âge Rupélien (Oligocène inférieur
belge). Ce fascicule est consacré à l’examen des tubes de térédiniformes présents dans la plupart de ces
fragments de bois pétrifiés.
Mots-clés: Belgique, Oligocène, Rupélien, Sables de Ruisbroek, Niveau à Phosphorites de Sint-Niklaas,
Teredinidae, Xylophagaidae, Terediniformes, Myoidea, Anomalodesmata, Bivalvia, Mollusca, Invertebrata.
Samenvatting
(Hilde Van Waes)
Bij het eerste onderzoek van de fossielen die opgeslagen waren in de schuif: SVK - Vegetal Remains, was het een
grote verrassing vast te stellen dat enkele gesilificiëerde houtstukken tubes van Teredinidae (Bivalvia, Mollusca)
bevatten, andere sporen van activiteiten van larvae van hout etende insecten (Coleoptera, Insecta) en dat de elf
lange sideriet concreties fragmenten van tubes van sabellid-achtige dieren (Sabellida, Annelida) zouden kunnen
zijn.
Deze Géominpal Belgica is het tweede supplement bij de Publicatie* die het onderzoek van de resten van
Invertebrata gevonden in het Sint-Niklaas Fosforiet Bed van Rupeliaan Ouderdom (Belgisch Onder Oligoceen)
tot doel had. Dit fasciculum is gewijd aan het onderzoek van de teredinid-tuben, die in het merendeel van deze
versteende houtfragmenten aanwezig zijn.
Sleutelwoorden: België, Oligoceen, Rupeliaan, Sint-Niklaas Phosphorite Bed, Teredinidae, Xylophagaidae,
Terediniformes, Myoidea, Anomalodesmata, Bivalvia, Mollusca, Invertebrata.
Kurzfassung
(Detlev Thies)
Eine erste Untersuchung der Fossilien in der Schublade kommt zu folgendem Ergebnis: SVK – Pflanzenreste.
Einige silifizierte Holzreste werden von Bohrmuschel-Röhren (Family Teredinidae, Bivalvia, Mollusca)
durchzo- zogen. Fraßspuren gehen auf Holz fressende Käfer (Coleoptera, Insecta) zurück. Bei elf länglichen
Siderit-Konkretionen handelt es sich möglicherweise um Fragmente der Ausfüllungen von Wohnröhren sabellidartiger Anneliden (Sabellida, Polychaeta, Annelida).
Diese Ausgabe von Géominpal Belgica ist die zweite Ergänzung zu Géominpal Belgica. 5(2)*, die sich mit der
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Untersuchung der Invertebraten-Reste aus dem Rüpel zeitlichen Sint-Niklaas Phosphorit Bed (Belgien, UnterOligozän) beschäftigt. Diese Ausgabe behandelt die Wohnröhren von Bohrmuscheln (Family Teredinidae), die
in der Mehrzahl der fossilen Holzreste vorhanden sind.
Schlüsselworten: Belgien, Oligozän, Rupelium, Sande von Ruisbroek, Sint-Niklaas Phosphorite Bed, Teredinidae, Xylophagaidae, Terediniformes, Myoidea, Anomalodesmata, Bivalvia, Mollusca, Invertebrata.
Resumen
(Jacques Herman)
Investigando por primera vez los fósiles en el cajón SVK - Vegetal Remains, tuvimos la sorpresa que unas
de las piezas de madera silicificada contuvieran tubos de Teredinidae (Bivalvia, Mollusca) y otras, huellas
de actividades de larvas de insectos xilófagos (Coleoptera, Insecta). Además once concreciones en siderita
podrían ser fragmentos de tubos de sabellid animales (Sabellida, Polychaeta, Annelida).
Este Géominpal Belgica es el segundo suplemento de la Publicación* dedicada a la investigación sobre los
restos de los invertebrados encontrados en el Horizonte con Fosforitas de Sint-Niklaas de la época
del Rupeliense. Este fascículo es dedicado al examen de los teredinid tubos presentes en la mayoría de
estos fragmentos de madera petrificados.
Palabras clave: Bélgica, Oligoceno, Rupeliense, Arenas de Ruisbroek, Sint-Niklaas Phosphorite Bed, Teredinidae, Xylophagaidae, Terediniformes, Myoidea, Anomalodesmata, Bivalvia, Mollusca, Invertebrata.
Resumo
(Jacques Herman)
Examinando pele primeira vez a gaveta ‘Restes végétaux’, grande era a surpresa para notar que certos pedaços de
madeira contiveram tubos de Teredinidae (Bivalvia, Mollusca), outros rastos de atividades de larvas de insetos
xylophagous (Coleoptera, Insecta) e onze fragmenentos de concreções sideriticos de forma muito esticada de
sabellid animais (Sabellida, Polychaeta, Annelida).
Este Géominpal Belgica estabelece a segunda adição na publicação* tendo tido para tema o estudo dos restos dos
Invertebrados descobriram dentro nele Horizon com Phosphorites de Sint-Niklaas de Rupélien (belga abaixe
Oligoceno). Esta parte é consagrada ao exame dos teredinid tubos estão presentes na maioria dos fragmentos de
madeira petrificada.
Palavras chaves: Bélgica, Oligoceno, Rupeliense, Areias de Ruisbroek, Horizon com Phosphorites de Sint-Niklaas, Teredinidae, Xylophagaidae, Terediniformes, Myoidea, Anomalodesmata, Bivalvia, Mollusca, Invertebrata.
Резюме
(Evgeny Popov)
Первое изучение ископаемых из ящика «SVK – растительные остатки» позволило с удивлением
обнаружить, что некоторые из окремнелых кусков древесины содержат трубки моллюсков-древоточцев
Teredinidae (Bivalvia, Molluska), другие – являются следами активности личинок насекомых-ксилофагов
(Coleoptera, Insecta), и что двенадцать удлиненных сидеритовых конкреций могут быть фрагментами
трубок сабелиидоподобных животных (Sabellida, Polychaeta, Annelida).
Этот выпуск Géominpal Belgica является вторым дополнением к публикации*, сфокусированной на изучении остатков беспозвоночных, которые были обнаружены в фосфоритовом горизонте Синт Никлаас
рюпельского возраста (нижний олигоцен Бельгии). Этот выпуск посвящен экспертизе трубок терeдинид,
встреченных в большом количестве во фрагментах окаменевшей древесины.
Ключевые слова: Бельгия, олигоцен, рюпель, пески Руисбрек, фосфоритовый горизонт Синт Никлаас,
беспозвоночные, Teredinidae, Xylophagaidae, Terediniformes, Myoidea, Anomalodesmata, Bivalvia, Mollusca, Invertebrata.
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2. Introduction
Discoveries of wood pieces bored by teredinid bivalves are relatively common in all the marine deposits of the
Belgian Neogene. But, in these Formations, the wood fragments are not petrified.
The presence of tubes of teredinid-like marine bivalves in these petrified pieces of wood implies that these
drifted pieces of wood were petrified after a silicifying * phase in an epi-continental environment.
*Induration, or petrification, is more correct because no chemical analysis certifies that it was a silicifying process.
Later, these petrified pieces of wood were dis-embedded from their original level, or levels, and concentrated in
the Sint-Niklaas Phosphorite Bed.
It is the reconstitution of the precise succession of all these events which must allow a better understanding of
the climatic variations of this period and the taphonomic complexity of this Horizon.
The determination, even at the generic level, of these tube-builder bivalves is confronted with a complex and
hazardous problematic. The zoological literature virtually furnishes no indications about diverse data which
could help paleontologists. Some examples are:
Is the cohabitation of diverse species of xylophagous molluscs in the same piece of wood conceivable? Do these
xylophagous molluscs have selective vegetal host(s) or not? Is the form of the section of their tubes a valid
criterion? Is a continuous rectilinear growth of their tubes a valid criterion?
The senior-author admits that all these data could allow proposing a plausible generic determination of these
teredinid remains, but without being absolutely certain that his proposal is correct.
3. Introduction
Les découvertes de pièces de bois perforées par des bivalves térédiniformes sont relativement communes dans
tous les dépôts marins du Néogène belge. Mais, dans ces Formations, les fragments de bois ne sont pas pétrifiés.
La présence de tubes de mollusques marins térédiniformes dans des fragments de bois pétrifiés implique que ces
pièces de bois flottées ont été indurées après avoir été perforées par ces invertébrés et qu’elles ont subi une
silicification* en milieu épicontinental.
*Induration, ou pétrification, est plus correct car aucune analyse chimique ne garantit qu’il s’agisse d’un processus de
silicification.
Ces pièces de bois indurées furent ensuite extraites d’une, ou de plusieurs strates, et reconcentrées dans
l’Horizon à Phosphorites de Sint-Niklaas.
C’est la reconstitution de l’ordre de succession de tous ces évènements qui devrait permettre d’essayer de mieux
comprendre les variations climatiques de cette période et la complexité taphonomique de cet Horizon.
La détermination, même au niveau générique, des bivalves qui ont construit ces tubes se heurte à une problématique complexe et hasardeuse. La littérature zoologique reste pratiquement muette en ce qui concerne des facteurs
qui pourraient venir en aide aux paléontologistes. En voici quelques exemples.
La cohabitation de diverses espèces de mollusques xylophages au sein d’une même pièce de bois est-elle concevable ? Ces xylophages s’attaquent-ils à tout bois flottant, ou à un groupe végétal précis ? La constance du diamètre de la section de leurs tubes est-t-elle un critère utile ? Ou encre, une croissance linéaire continue est-elle
un indice ?
Le senior-auteur admet que toutes ces données devraient permettre de proposer une détermination générique
plausible de ces restes de térédiniformes, sans pour autant prétendre que celle proposée soit la bonne.
Introductie
Vondsten van houtfragmenten geboord door teredinid-achtige tweekleppige mollusken zijn relatief frequent in al
de mariene afzettingen van het Belgische Neogeen. Maar, in deze Formaties, zijn de houtfragmenten niet
versteend.
De aanwezigheid van tuben van teredinid-achtige mariene tweekleppige mollusken in deze versteende hout frag-
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menten betekent dat deze drijvende houtfragmenten versteenden na een silicificatie* fase in een epi-continentale
omgeving.
*Induratie, of petrificatie, is correcter omdat geen enkele chemische analyse garandeert dat het een silicificatie proces was.
Later kwamen deze versteende houtfragmenten los uit hun originele laag, of lagen, en werden ze in het SintNiklaas Phosphorite Bed geconcentreerd.
Het is de reconstitutie van de precieze successieve fenomenen die een beter begrip moet geven aan de
klimatologische variaties van deze periode en aan de tafonomische complexiteit van dit Horizon.
De determinatie, zelfs op niveau van het Genus, van deze tweekleppige tuben-makers is geconfronteerd met een
complexe en dubieuze problematiek. De zoölogische literatuur geeft geen enkele indicatie over diverse gegevens
die voor de paleontologen van groot belang kunnen zijn. Enkele voorbeelden zijn:
Is het samenleven van diverse soorten van hout-etende mollusken in het zelfde houtfragment mogelijk? Kiezen
deze hout-etende mollusken een bepaalde plantensoort of niet? Is de vorm van de sectie van hun tuben een
betrouwbaar criterium? Is een constante en rechtlijnige groei van hun tuben een betrouwbaar criterium?
De senior-auteur denkt dat al deze gegevens een geloofwaardige determinatie van deze teredinid fossielen
mogelijk zou moeten maken, maar blijft twijfelen of zijn voorstel correct is.
5. Phylum Mollusca LINNAEUS, 1758
5.1. Systematics
The Phylum Mollusca regroups the seven following Classes:
Class Bivalvia LINNAEUS, 1758, Class Cephalopoda CUVIER, 1797,
Class Polyplacophora de BLAINVILLE, 1816, Class Scaphopoda BRONN, 1862,
Class Solenogastres GEGENBAUR, 1878, Class Gastropoda CUVIER, 1895,
and Class Monoplacophora ORDNER, 1940.
5.2. Remarks
In some Classifications, Aplacophora is used for Scolenogastres. Except for the Solenogastres, all these Classes
are represented by extant and extinct taxa.
Searching for recent systematics of diverse Families and Genera of Bivalvia, the senior-author has compared the
more recent lists proposed by ADW, ITIS, MarBEF, Taxonomicom or WoRMS. Constating the huge difference
of the interpretation of these systematic ranks proposed by some of these lists, the senior-author has choised a
mixed interpretation for which he is the sole responsible.
Therefore, in case of doubt, the senior-author use the conditional term, such as: This Genus could regroups the
following extant specific taxa. WoRMS 2015 is in fact WoRMS 2013, but nothing was modified during these
two years.
6. Class Bivalvia
6.1. Generalities
The Class Bivalvia regroups a little more than a hundred Families, but only two are only represented by woodborers taxa.
Their first records date from the Lower Cambrian Period*, they regroup more than 1200 Genera which include
more than 10.000 living species inhabiting marine, brackish or fresh waters as well as all the continental
environments.
*Circa 510 million years ago.
6.2. Systematics
Only two Families of the Order Myoida STOLICKA, 1870 (Class Bivalvia) are represented by
strictly xylophagous* taxa:
The Family Teredinidae RAFINESQUE, 1815 and the Family Xylophagaidae HAGA & KASE, 2013.
*Which means that all their extant representatives and, so far as known,
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all their extinct representatives feed, or fed, on vegetal remains.
These two Families are included in the Order Myoida STOLICKA, 1870 which only comprises
the four Families:
Family Myidae de LAMARCK, 1809, Family Pholadidae de LAMARCK, 1809,
Family Corbulidae de LAMARCK, 1818 and
Family Gastrochaenidae GRAY, 1840.
Systematic remark
Some malacogists consider that the Family Hiatellidae GRAY, 1824 may be included in this Order.
See also the Chapter 11 which details the recent proposal of the Order Anomalodesmata.
References
BIELER, R., CARTER, J. G. & COAN, E. V. 2010 : Classification of Bivalve families. Pp.: 113-133 In:
BOUCHET, P. & ROCROI, J.-P. 2010: Nomenclator of Bivalve Families. Malacologia. 52(2): 1-184.
Illustration
One species of the type-Genus of these five Families is illustrated on the diverse comparison Plates.
Comparison
A maximum of the extant and extinct taxa of the Families Teredinidae and Xylophagaidae
are examined in this Publication in accordance with their more recent classification.
7. Family Teredinidae RAFINESQUE, 1815
7.1. Systematics
According to ITIS 2015 and WoRMS 2015, this Family regroups three Sub-Families:
Sub-Family Teredininae RAFINESQUE, 1815, Sub-Family Kuphinae TRYON, 1862
and Sub-Family Bankiinae TURNER, 1966.
All together they are represented by some fifteen extinct and extant Genera.
This list is presented in function of the date on which each of these Sub-Families was proposed.
7.2. Sub-Family Teredininae RAFINESQUE, 1815
(Plates 9, 10, 15 and 19)
Systematics
According to ITIS 2015 and WoRMS 2015, this Sub-Family includes the ten extant Genera:
Genus Teredo LINNAEUS, 1758, Genus Uperotus GUETTARD, 1770, Genus Lyrodus GOULD, 1870,
Genus Bactronophorus TAPPARONE CANEFRY, 1877, Genus Neoteredo BARTSCH, 1920,
Genus Teredora BARTSCH, 1921, Genus Teredothyra BARTSCH, 1921, Genus Psiloteredo BARTSCH, 1922,
Genus Dicyathifer IREDALE, 1932 and Genus Zachsia BULATOFF & RJABSCHIKOFF, 1933.
This list is presented in function of the date on which each of these Genera was proposed.
Paleontological remark
The Genus Teredina is a Genus proposed by de Lamarck in 1818 and based on the taxon Teredina personata
(de LAMARCK, 1806) in which plenty of teredinid-like records encountered in European, Asian and northern
American strata of Upper Cretaceous to Pliocene Ages were regrouped.
The frequent lack of distinctive arguments that allow distinguishing these taxa from the extant taxa of the Family
Teredinidae obliges the scientific community to consider some species attributed to this Genus as doubtful taxa.
Singularities
All the extant representatives of this Family possess a bivalve shell having lost its primordial function: the
protection of the body of their owner. Their shell has evolved to become a perfect boring instrument of drifting
pieces of wood and human wooden naval constructions.
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Their galleries are protected by a calcareous tube of variable thickness, size and morphology. The extern side of
their valves present small extern asperities oriented towards the wooden walls.
By the combination of three factors: the suction their foot realises, the contraction of their retractor muscles and
the swelling of the anterior part of their body, these molluscs bore their individual gallery.
The alternative contractions and retractions of their two adductor muscles induce a permanent rotation of their
valves along a central axis, which helps to rasp the piece of wood.
The anterior part of their body is also able to turn regularly left and right, giving a perfect circular section to their
galleries of which the posterior aperture may be closed by two bio-mineralized pallets.
Such a sophisticated adaptation theoretically requires a very long time of evolution and important modifications
of the genetic code.
The modifications of the genetic code of the Teredinidae are responsible for the extreme reduction of the size * of
their shell, the sudden passage from a filtering feeding mode to a strict xylophagous feeding mode, the
construction of a very long calcareous tube protecting their hyper-elongated siphon and the formation of one pair
of bio-mineralised appendices called pallets.
*The length of shells of Myoida rises easily up to ten centimetres. Teredinid-shells are commonly millimetric-sized.
Life cycle and feeding mode
Life cycle
This one may be summarized as follows: Egg - Larval drifting stage - Fixation on a wooden drifting support –
Penetration into its support - Boring activity - Death.
Feeding mode
Contrarily to the extinct and extant representatives of the Family Pholadidae de L AMATCK, 1809 which bore
vertical holes in organic or inorganic calcareous supports, all the members of the Family Teredinidae bore
galleries which, after penetration into the wood, follow the structure of the wood they parasite.
It seems that the ability to digest the cellulose of the wood of all the extant representatives of the Teredinidae
only results from their symbiotic association with Bacteria, such as Teredinibacter turnerae DISTEL, MORRILL,
MacLAREN-TOUSSAINT, FRANKS & WATERBURY, 2002.
Growth
The growth of the tubes of the diverse taxa of the extant representatives of this Family seems to be a resultant of
diverse factors such as the hardness of the wood they bore and the salinity of the waters. For some African
species, their size may increase from two to three centimetres in one month.
Ecology
All the extant populations of the Family Teredinidae feed upon pieces of wood
drifting in marine or brackish waters.
1. Genus Teredo LINNAEUS, 1758
(Plates 9, 10 and 15)
Systematics
According to ITIS 2015, the Genus Teredo is based on Teredo navalis LINNAEUS, 1758
and could regroup the thirteen other species: Teredo modica DESHAYES, 1856,
Teredo furcifera MARTENS in SEMON, 1894, Teredo clappi BARTSCH, 1923, Teredo mindanensis BARTSCH, 1923,
Teredo bartschi CLAPP, 1923, Teredo fulleri CLAPP, 1924, Teredo johnsoni CLAPP, 1924,
Teredo portoricensis CLAPP, 1924, Teredo somersi CLAPP, 1924, Teredo poculifer IREDALE, 1936,
Teredo aegyptos MOLL, 1941, Teredo triangularis EDMONSON, 1942 and Teredo bitubula LI, 1965.
Ecology and Geographical distribution
All the extant populations of the Genus Teredo feed upon pieces of wood drifting in marine or brackish waters
and all their individuals produce a protective calcareous tube.
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The Genus itself has a world-wide distribution, but some populations of its representatives, such as T.
mindanensis and T. portoricensis have, so far as known, restricted areas of distribution.
Geological record
From the Albian Stage (Upper Cretaceous) to the Present Times.
References for Genus Teredo
COE, W. R. 1933: Sexual phases in Teredo. Biological Bulletin. 65: 283-303.
GRAVE, B. H. 1928: Natural history of shipworm, Teredo navalis, at Woods Hole, Massachusetts. Biological
Bulletin. 55: 260-282.
MILLER, R. C. 1924: The boring mechanism of Teredo. University of California Publications in Zoology. 26:
41-80.
THOMPSON,W. 1847: Note on the Teredo norvegica (T. navalis, Turton, not Linn.), Xylophaga dorsalis,
Limnoria terebrans and Cheluda terebrans, combined in destroying the submerged wood-work at the harbor of
Ardrossan on the coast of Ayrshire. Annals and Magazine of Natural History. 20: 157-164.
TURNER, R. D. 1966: A survey and illustrated catalogue of the Teredinidae (Mollusca: Bivalvia). The Museum
of Comparative Zoology, Harvard University, Cambridge. 265 p.
2. Genus Uperotus GUETTARD, 1770
(Plate 18)
Systematics
According to ITIS 2015, the Genus Uperotus only regroups two extant species:
Uperotus clavus (GMELIN, 1791) and Uperotus panamensis (BARTSCH, 1922).
Fossil record
None, except if the Belgian Eocene colonies discovered in the Belgian Lutetian
could be attributed to this Genus.
Geographical distribution
The extant representatives of this Genus are inhabitants of the Eastern Pacific Ocean:
Uperotus clavus seems to be endemic to some waters of New Zealand and
Uperotus panamensis is an inhabitant of the western tropical waters of America.
References for the Genus Uperotus
MYRA, A. 1971: Sea Shells of Tropical West America: Marine Mollusks from Baja California to Peru. See p.:
281.
OLIVER, W., R., B. 1915: The Mollusca of the Kermadec Islands. Transactions and Proceedings of the New
Zealand Institute. 47: See p.: 556.
3. Genus Lyrodus GOULD, 1870
Systematics
According to the researches of the senior-author, the Genus Lyrodus is represented by its generotype:
Lyrodus pedicellatus (de QUATREFAGES, 1849)
and, at least, by the four following extant species:
Lyrodus bipartitus (JEFFREYS, 1860), Lyrodus floridanus (BARTSCH, 1922),
Lyrodus takanoshimensis (ROCH, 1929) and Lyrodus medilobata (EDMONSON, 1942).
Ecology and distribution
The populations of these taxa are scattered
off the coasts of Florida (U.S.A.), off these of New Zealand,
off these of eastern Australia and in the Coral Sea.
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Geological record
Taxa attributed to this Genus are reported from strata of Upper Cretaceous Age to the Present Times.
References for Genus Lyrodus
CALLOWAY, C. B. & TURNER, R. D. 1983: Documentation and implications of rapid successive gametogenic
cycles and broods in the shipworm Lyrodus floridanus (Bartsch) (Bivalvia, Teredinidae). Journal of Research
Shellfish. 3: 65-69.
COAN, E. V. & VALENTICH-SCOTT, P. 2012: Bivalve seashells of tropical West America. Marine bivalve
mollusks from Baja California to northern Peru. 2 vols., 1258 p.
DISTEL, D. L., BEAUDOIN, D. J. & MORRILL, W. 2002: Coexistence of multiple proteobacterial
endosymbionts in the gills of the wood-boring bivalve Lyrodus pedicellatus (Bivalvia: Teredinidae). Applied and
Environmental Microbiology. 68: 6292-6299.
LUYTEN, Y. A., THOMPSON, J. R., MORRILL, W., POLZ, M. F. & DISTEL, D. L. 2006: Extensive variation
in intracellular symbiont community composition among members of a single population of the wood-boring
bivalve Lyrodus pedicellatus (Bivalvia:Teredinidae). Applied Environmental Microbiology. 72: 412-17.
TURNER, R. D. 1966: A Survey and Illustrated Catalogue of Teredinidae (Mollusca: Bivalvia). Cambridge.
Massachusetts: Museum of Comparative Zoology. 265 p. See p.: 78.
4. Genus Bactronophorus TAPPARONE CANEFRY, 1877
Systematics
According to ITIS 2015, the Genus Bactronophorus seems to be represented
by its generotype and single species Bactronophorus thoracites (GOULD, 1856).
Its habitat was described in 1856 as the mud of mangrove swamps in the torrid South West Pacific
and considered as a mobile boring herbivore by Gould, his inventor.
Its pallets are made of phosphate calcium carbonate.
Geological record
None.
CALLOWAY, C. B. & TURNER, R. D. 1983: Documentation and implications of rapid successive gametogenic
cycles and broods in the shipworm Lyrodus floridanus (Bartsch) (Bivalvia, Teredinidae). Journal of Shellfish
Research. 3: 65-69.
JEFFREYS, J. G. 1860: A synoptical list of the British species of Teredo with a notice of the exotic species.
Annals and Magazine of Natural History. (3)6: 121-127. See p.: 123.
MACINTOSH, H. 2012: Lyrodus turnerae, a new teredinid from eastern Australia and the Coral Sea (Bivalvia:
Teredinidae). Molluscan Research. 32(1): 36-42.
MOLNAR, J. L., GAMBOA, R., REVENGA, C. & SPALDING, M. 2008: Assessing the global threat of invasive species to marine biodiversity: Framing the big picture. Frontiers in Ecology and the Environment. 6(9): 485492.
QUATREFAGES, (de) A. 1849 : Mémoire sur le genre Taret (Teredo Linn.). Annales des Sciences Naturelles.
(Zoologie Biologie Animale). 11: 19-64, pls.: 1-5.
TURNER, R. D. 1966: A Survey and Illustrated Catalogue of Teredinidae (Mollusca: Bivalvia). Cambridge
(Massachusetts): Museum of Comparative Zoology. 265 p. See p.: 86 and p.: 101.
5. Genus Neoteredo BARTSCH, 1920
Systematics
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This monospecific Genus is based on Neoteredo reynei BARTSCH, 1920.
The rare populations identified with certitude inhabit some tropical brackish * water areas of the western Atlantic
coasts and bore stem and adventive roots of diverse extant taxa of the Family Rhizophoraceae P ERSOON, 1806
(Angiospermae).
*As usual, in place of brackish species, WoRMS mentions: marine species.
As far as known, populations of this species were only observed
in some tropical mangrove areas of the western Atlantic coasts.
Geological record
None.
ALVES, R. R. N. & DIAS, T. L. P. 2010: Usos de invertebrados na medicina popular no Brasil e suas implicacoes para conservaçao. Tropical Conservation Science. 3(2): 159-174. See p.: 171.
AVIZ, D., FERREIRA de MELLO, C. & FERNANDES da SILVA, P. 2009: Macrofauna associada as galerias
de Neoteredo reynei (Bartsch, 1920) (Mollusca: Bivalvia) em troncos de Rhizophora mangle Linnaeus durante o
período menos chuvoso, em manguezal de Sao Caetano de Odivelas, Para (costa norte do Brasil). Boletim do
Museu Emílio Goeldi. Ciencias Naturais. 4(1): 47-55.
DE MORAES, D, T. & LOPES, S. G. B. 2003: Neoteredo reynei (Bartsch, 1920) (Bivalvia, Teredinidae). Journal of Molluscan Studies. 69(4): 311-318.
FILHO, C. S., TAGLIARO, C. H. & BEASLEY, C. R. 2008: Seasonal abundance of the shipworm Neoteredo
reynei (Bivalvia, Teredinidae) in mangrove driftwood from a northern Brazilian beach. Iheringia. Serie
Zoologia. 98: 17-23.
TURNER, R. D. 1966: A Survey and Illustrated Catalogue of Teredinidae (Mollusca: Bivalvia). Cambridge (Massachusetts). Museum of Comparative Zoology. 265 p. See p.: 73 and p.: 113.
6. Genus Teredora BARTSCH, 1921
Systematics and Distribution
The Genus Teredora seems to regroup the two sole extant species:
Teredora malleolus (TURTON, 1822), living in the Caribbean Sea
and Teredora princesae (SIVICKIS, 1928), living in the western Pacific Ocean.
Ecology
Both species live in pieces of wood drifting in marine waters.
Geological record
None.
References for Genus Teredora
KILBURN, R. N. & RIPPEY, E. 1982: Sea Shells of Southern Africa. Macmillan South Africa. Johannesburg.
249 p. See p.: 205.
LOCARD, A. 1886: Prodrome de malacologie française. Catalogue général des mollusques vivants de France.
Mollusque marins. Lyon, H. Georg & Paris, Baillière. X + 778 p.
STEYN, D. G. & LUSSI, M. 1998: Marine Shells of South Africa. An Illustrated Collector’s Guide to Beached
Shells. Ekogilde Publishers, Hartebeespoort, South Africa, II + 264 p. See p.: 248.
TURNER, R. D. 1966: A Survey and Illustrated Catalogue of Teredinidae (Mollusca: Bivalvia). Cambridge (Ma
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-ssachusetts): Museum of Comparative Zoology. 265 p. See p.: 75.
TURTON, W. 1822: Conchylia dithyra Insularum britannicarum. The bivalve shells of the British Islands. M.A.
Natali, London, & Combe and son, Leicester. XLVII + 279 p., 20 pls.
7. Genus Teredothyra BARTSCH, 1921
Systematics
The Genus Teredothyra could regroup the five extant species:
Teredothyra excavata (JEFFREYS, 1860), Teredothyra dominicensis BARTSCH, 1921,
Teredothyra matocotana BARTSCH, 1927, Teredothyra smithi BARTSCH, 1927
and Teredothyra remiformis LI, 1965.
Details
Shells of some populations of Teredothyra matocotana were collected in stranded pieces of wood along the
western Australian coasts. Teredothyra smithi and Teredothyra remiformis are reported as fairly common in the
Chinese waters. Presence of Teredothyra dominicensis are only reported from Caribbean waters and Teredothyra
smithi seems to be a common inhabitant of the tropical waters of the western Pacific.
Conclusion
This Genus has a world-wide distribution, but its extant representatives inhabit specific or con-specific zones.
Geological record
None
References for Genus Teredothyra
Annals and Magazine of Natural History. (3)6: 121-127.
8. Genus Psiloteredo BARTSCH, 1922
Systematics
According to ADW 2014, the Genus Psiloteredo regroups the three following species:
Psiloteredo dilitata (STIMPSON, 1851), Psiloteredo megotara (HANLEY in FORBES & HANLEY, 1848)
and Psiloteredo nana (TURTON, 1822), all previously attributed to the Genus Teredo.
Eastern North Atlantic.
Geological record
None.
References for Genus Psiloteredo
DEFRANCE, J. L. M. 1804-1845: Dictionnaire des Sciences Naturelles dans lequel on traite méthodiquement
des différents êtres de la nature. Paris Vol. 32. See p.: 361.
(Massachusetts): Museum of Comparative Zoology. 265 p. See p.: 76.
9. Genus Dicyathifer IREDALE, 1932
Systematics
According to ITIS 2015 and WoRMS 2015,
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the Genus Dicyathifer is based on its single extant representative: D. mannii (WRIGHT, 1866).
D. mannii is commonly found in mangrove environments on the coasts of the Arabian Sea.
Geological record
None.
References for Genus Dicyathifer
DEY, A. 2006: Handbook on Mangrove Associate Molluscs of Sundarbans. XII + 96 p.
10. Genus Zachsia BULATOFF & RJABSCHIKOFF, 1933
Systematics
This monospecific Genus is based on:
Zachsia zenkewitschi BULATOFF & RJABSCHIKOFF, 1933.
According to WoRMS 2015,
the first representative individuals of Zachsia zenkewitschi
were discovered in a marine environment.
This species seems to have a world-wide distribution, in icy to tropical waters.
Geological record
None.
Reference for Genus Zachsia
7.3. Sub-Family Kuphinae TRYON, 1862
Systematics
This mono-generic Sub-Family is based on its single extant representative:
Genus Kuphus GUETTARD, 1770.
Genus Kuphus GUETTARD, 1770
Systematics
According to WoRMS 2015,
the Genus Kuphus regroups three taxa,
two extinct ones:
Kuphus incrassatus GABB, 1873 and Kuphus fistula LEA, 1843,
and one extant species:
Kuphus polythalamia (LINNAEUS, 1758).
The diverse populations of the extant species, Kuphus polythalamia, may be encountered in various parts of the
western Pacific Ocean, the eastern Indian Ocean and the Indo-Malaysian area, including the Philippines, Sumatra
and Mozambique waters.
The majority of the fossils attributed to the Genus Kuphus require revision. Diverse fossils Kuphus are in fact*
tubes of carinate serpulid worms (Family Serpulidae JOHNSTON, 1865, Polychaeta, Annelida).
*See the specimen illustrated as: Image for Genus Kuphus on www.wikipedia.org.
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Fossils attributed to Kuphus polythalamia have been found in various sediments * of Oligocene Age and representatives of tropical and sub-tropical environments.
*In Indonesia, Pakistan, Jamaica, Grenada, South Africa and Somalia.
Fossils attributed to Kuphus incrassatus have been reported from sediments of Oligocene and Miocene Ages
in Jamaica, Mexico, Panama, Puerto Rico, Trinidad, Tobago, Florida (U.S.A.) and Mississippi (U.S.A.).
They were also used for absolute dating of these rocks, using the relative proportions of two strontium isotopes.
Fossils attributed to Kuphus fistula of Miocene and Pliocene Ages have been reported from diverse localities in
Virginia (U.S. A.).
Ecology and Geographical distribution
The environment where the extant populations of the Genus Kuphus live is nearly similar to this where extant
populations of the Genus Teredo live. All their extant individuals produce a protective calcareous tube.
Ancestors
Such as suggested by Ruth Turner in 1966, it is not sure that their ancestors were able to bore galleries into wood
and maybe they simply lived embedded in soft sediments.
References for Genus Kuphus
DAUTZENBERG, P. 1929 : Contribution à l'étude de la faune de Madagascar: Mollusca marina testacea.
Faune des colonies françaises. III (fasc. 4). Société d'Editions géographiques, maritimes et coloniales: Paris.321636, pls.: IV-VII.
(Massachusetts): Museum of Comparative Zoology. IX + 265 p. See p.: 7.
7.4. Sub-Family Bankiinae TURNER, 1966
Systematics
This Sub-Family includes the four extant Genera:
Genus Bankia GRAY, 1842, Genus Nausitora WRIGHT, 1864,
Genus Nototeredo BARTSCH, 1923 and Genus Spathoteredo MOLL, 1928.
1. Genus Bankia GRAY, 1842
(Plates 11 and 19)
Systematics
A comparison between two official lists seems obligatory.
According to WoRMS
The Genus Bankia is based on Bankia carinata (GRAY, 1827) and regroups
the twenty-three other extant species:
B. bipalmulata (de LAMARCK, 1801), B. bipennata (TURTON, 1819), B. carinata (GRAY, 1827),
B. setacea (TRYON, 1860), B. brevis (DESHAYES, 1863), B. martensi (STEMPELL, 1899),
B. gouldi (BARTSCH, 1908), B. australis (CALMAN, 1920), B. zeteki BARTSCH, 1921, B. orcutti BARTSCH, 1923,
B. barthelowi BARTSCH, 1927, B. philippinensis BARTSCH, 1927, B. anechoensis ROCH, 1929,
B. campanellata MOLL & ROCH, 1931, B. fimbriatula MOLL & ROCH, 1931, B. rochi MOLL, 1931,
B. nordi MOLL, 1935, B. cieba CLENCH & TURNER, 1946, B. destructa CLENCH & TURNER, 1946,
B. forsteri CLENCH & TURNER, 1946, B. gracilis MOLL, 1935, B. insularis MUNARI, 1979 and
B. netzalia TURNER & MCKOY, 1979.
As well as the three extinct taxa:
B. turneri POWELL & BARTRUM, 1929, B. aemula LAWS, 1944 and B. cupedia LAWS, 1944.
According to ITIS
The Genus Bankia is based on Bankia carinata (GRAY, 1827) and includes the thirteen other species:
B. bipennata (TURTON, 1819), B. brevis (DESHAYES, 1863), B. setacea (TRYON, 1863),
B. martensi (STEMPELL, 1899), B. gouldi (BARTSCH, 1908), B. australis (CALMAN, 1920),
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B. zeteki (BARTSCH, 1921), B. fimbriata MOLL & ROCH, 1931, B. sibirica ROCH, 1934,
B. cieba CLENCH & TURNER, 1946, B. destructa CLENCH & TURNER, 1946,
B. fosteri CLENCH & TURNER, 1946 and B. neztalia (TURNER & McKOY, 1979).
Remark concerning the enormous difference between these two lists
The responsible ones for ITIS mention only the extant taxa, but have taken in consideration the numerous
publications which allowed reducing the number of the extant taxa from twenty-four to fourteen and the
responsible ones for WoRMS simply ignore all these scientific works.
All the representatives of the Genus Bankia are common drifting wood feeders. The principal zone of
distribution of this Genus seems to be the West Indies. Its occurrence in the North Sea could result from the
formation of the Gulf Stream.
Geological record
Fossil taxa attributedof this Genus have been reported from the early American Cretaceous.
References for Genus Bankia
mollusks from Baja California to northern Peru. 2 vols., 1258 p.
COTTON, B. C. 1934: Pelecypoda of the Flindersian region, southern Australia. No. 3. Records of the South
Australian Museum. 5(2): 173-178.
SIPE, A. R., WILBUR, A. E. & CARY, S. C. 2000: Bacterial symbiont transmission in the wood-boring
shipworm Bankia setacea (Bivalvia:Teredinidae). Applied Environmental Microbiology. 66: 1685-1691.
TURNER, R. D. 1955: The family Pholadidae in the western Atlantic and the eastern Pacific, Part II:
Martesiinae, Jouannetiinae and Xylophagainae. Johnsonia. 3: 65-160.
(Massachusetts). Museum of Comparative Zoology. 265 p. See p.: 80.
TURNER, R. D. & MCKOY, J. L. 1979: Bankia neztalia n. sp. (Bivalvia: Teredinidae) from Australia-New
Zealand, and its relationships. Journal of the Royal Society of New Zealand. 9: 465-473.
2. Genus Nausitoria WRIGHT, 1884
Systematics
According to WoRMS 2015, the Genus Nausitoria * was proposed for a fresh water Teredinidae discovered in
the Comer River, a branch of the Gange (India): Nausitoria fusticulus (JEFFREYS, 1860).
According to WoRMS 2015, the Genus Nausitoria * includes also the extant marine species: Nausitoria aurita
HEDLEY, 1899.
*WoRMS, alternatively opts for Nausitora or Nausitoria. Nausitoria is the good option !
If the second generic attribution is correct, this Genus is adapted to a large spectrum of s alinity.
Geological record
Taxa attributed to this Genus have been reported from strata of
Upper Cretaceous Age.
References for Genus Nausitoria
TURGEON, D. D., LYONS, W. G., MIKKELSEN, P., ROSENBERG, G. & MORETZSOHN, F. 2009:
Bivalvia (Mollusca) of the Gulf of Mexico. Pp.: 711-744 In: FELFER, D. L. & CAMP, D. K. Eds. 2009: Gulf of
Mexico-Origins, Waters, and Biota. Biodiversity. Texas A & M Press. See p.: 737.
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(Massachusetts). Museum of Comparative Zoology. 265 p. See p.: 93.
3. Genus Nototeredo BARTSCH, 1923
(Plate 11)
Systematics
According to ITIS 2014, the Genus Nototeredo is based on Nototeredo norvegica (SPENGLER, 1792)
and includes, at least, the two following other extant species:
Nototeredo edax (HEDLEY, 1895) and Nototeredo knoxi (BARTSCH, 1917).
Nototeredo edax is an Indo-Pacific species. Nototeredo norvegica is a common species inhabiting the north-east
Atlantic, from Iceland to the Mediterranean and Nototeredo knoxi is known by two western Atlantic
populations.
Morphology of the tubes of Nototeredo norvegica
(Plate 11)
It is the single extant taxon of the teredinid Bivalvia, the senior-author knows
that some of its individuals build tubes presenting successive different sized cross-sections.
Geological record
None.
References for Genus Nototeredo
mollusks from Baja California to northern Peru. 2 vols. 1258 p.
TURGEON, D. D., LYONS, W. G., MIKKELSEN, P., ROSENBERG, G. & MORETZSOHN, F. 2009:
Bivalvia (Mollusca) of the Gulf of Mexico. Pp.: 711-744. In: Felder, D. L. & Camp, D. K. Eds. 2009: Gulf of
Mexico-Origins, Waters and Biota. Biodiversity. Texas A & M Press.
(Massachusetts): Museum of Comparative Zoology. 265 p. See p.: 78.
4. Genus Spathoteredo MOLL, 1928
Systematics
According to the researches of the senior-author, the Genus Spathoteredo only regroups two species:
Spathoteredo spatha (JEFFREYS, 1860) and Spathoteredo obtusa (SIVICKIS, 1928).
All the populations of these two species were collected in mangrove environments
of the north-western Atlantic coasts of America.
Geological record
None.
References for Genus Spathtoteredo
GOFAS, S.; LE RENARD, J. & BOUCHET, P. 2001: Mollusca. In: COSTELLO, M. J., EMBLOW, C. & WHITE, R. J. Eds. 2001: European register of marine species: a check-list of the marine species in Europe and a
bibliography of guides to their identification. Collection Patrimoines Naturels. 50: 465 p. See: 180-213.
TURGEON, D. D., QUINN, J. F., BOGAN, A .E., COHAN, E. V., HOCHBERG, F. G., LYONS, W. G.,
MIKKELSEN, R. J., NEVES, C. F. E., ROPER, G., ROSENBERG, B., ROTH, A., SCHELTEMA, F. G.,
THOMPSON, M., VECCHIONE & WILLIAMS, J. D. & TURNER, R. D. 1998: Common and scientific names
of aquatic invertebrates from the United States and Canada: Mollusks 2d. Ed. American Fisheries Society.
Special Publication. 26: 526 p. See p.: 78.
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7.5. Geographical distribution of the Family Teredinidae
Principal data
The larvae of all the taxa attributed to this Family being nectic organisms, its world-wide distribution is easy to
understand, but the fact that many of its taxa only have restricted distribution zones remains unexplained.
Singularities
Some of its species are able to live in brackish water environments and even in fresh water environments, but the
ones that live in fresh water environments can no more tolerate marine waters.
7.6. Geological range of the Family Teredinidae
Problem of the validity of the Genus Teredina de LAMARCK, 1818
Systematics
According to Paleobiology Database, the Genus Teredina is based on the extinct European taxon: Teredina
personata (de LAMARCK, 1806) and includes also the Paleocene European taxa: Teredina oweni DESHAYES,
1856, and, at least, the three other extinct American taxa: Teredina fistula (LEA, 1843), Teredina laramiensis
(WHITEFIELD, 1902) and Teredina neomexicana STANTON, 1916.
Some tubes of Indonesian Teredinidae, discovered in Pliocene strata, were also attributed to this Genus.
Paleoecology
All the representative taxa of this extinct Genus are considered as feeders of drifting pieces of wood.
Geological record
From the Upper Cretaceous to the late Pliocene,
in Europe, Asia and North America.
Each member attributed to this extinct generic taxon requires a revision of its status.
References for Genus Teredina
GRAY, J. E. 1858: On the structure and position of the genus Teredina of Lamarck. Annals and Magazine of
Natural History. 3(2): 85-90.
GRAY, J., E. 1858: Further observations on the genus Teredina, Lamarck. Journal of Natural History. 3(3):
162-163.
LEA, H. C. 1843: Descriptions of Some New Fossil Shells, from the Tertiary of Petersburg, Virginia.
Philadelphia. 12 p.
LUDVIGSEN, R. & GRAHAM, B. 1997: West Coast Fossils: A Guide to the Ancient Life of Vancouver Island.
P.: 107.
TRYON (Jr.), G. W. 1863: Monograph of the Family Teredinidae. Proceedings of the Society for Natural Sciences of Philadelphia. 9: 453-482.
Extant north-western Atlantic associations of terediniformes
Leroux (1983) signalised that Lyrodus pedicellatus (de QUATREFAGES, 1849), Teredo navalis LINNAEUS, 1758
and Nototeredo norvegica (SPENGLER, 1792) cohabit in the Gulf of Morbihan (Bretagne, France).
7.7. Ichnology and the Teredinidae
Ichnologists have proposed diverse names to mention the discoveries of teredinid-like petrified structures, such
as Teredolithes HERRMANNSEN, 1852, Teredolites clavatus LEYMERIE,1842 and Teredolites longissimus
LEYMERIE,1842, Teredomorphus, HEINZE, 1943: Teredomorphus glaber (KRAATZ, 1895), Teredomorphus glaber
(KRAATZ, 1895), Teredomorphus rufipes (KRAATZ, 1895) and Genus Teredon NORTON, 1869: Teredon bartschi
CLAPP, 1936, Teredon cubensis CRESSON, 1865 and Teredon latitarsis CRESSON, 1865.
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All these denominations only point out the presence of teredinid-like invertebrates
in the strata in which they were discovered.
7.8. References for Family Teredinidae
ADERADO VIDAL, J. M. & ALMEIDA ROCHA-BARREIRA, (de), C. 2009: Teredos (Mollusca: Bivalvia:
Teredinidae) de um estuário do Nordeste brasileiro. Arquiyos de Ciencias de Mar. 42(2): 43-49.
ARDOVINI, R. & COSSIGNANI, T. 2004: Conchiglie dell'Africa Occidentale (incluse Azzorre, Madeira e Canarie). English-Italian edition. L' Informatore Piceno.Ancona, Italy. 319 p.
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(Bivalvia, Teredinidae). Journal of Molluscan Studies. 69: 311-318.
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McKOY, J. L. 1978: Records of Upper Cretaceous and Tertiary Bankia Gray (Mollusca: Teredinidae) from New
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NAIR, N. B. & GURUMANI, O. N. 1957 : A new shipworm Teredo (Teredora) vattanansis from the east coast
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OLIVER, P. G., WOOD, H. & HOLMES, A. M. 2009: First British record of the shipworm Uperotus (Bivalvia:
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ORTEGA-ARIZA, D. 2011: The utility of Kuphus incrassatus bivalves for determining absolute ages and
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POPHAM, J. D. & DIKSON, M. R. 1973: Bacterial associations in the teredo Bankia australis (Lamellibranchia: Mollusca). Marine Biology. 19: 338-340.
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POZARYSKA, K. & PUGACZEWESKA, H. 1981: Bivalve nature of Huene’s dinosaur Succinodon. Acta
Palaeontologica Polonica. 26(1): 27-34.
RANCUREL, P. 1955: Teredo thomsoni Tryon et Teredo lieberkindi Roch, transformations morphologiques des
palettes au cours de leur croissance. Bulletin de l’IFAN. XVIII(4): 8 pages. PDF on-line.
REISE, K., GOLLASCH, S. & WOLFF, W. J. 1999: Introduced marine species of the North Sea coasts.
Helgolander Meeresuntersuchungen. 52: 219-234.
ROCHA-BARREIRA, C. 2009: Teredos (Mollusca: Bivalvia: Teredinidae) de um estuário do Nordeste
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ROMANO, C., VOIGHT, J. R., PEREZ-PORTELA, R. & MARTIN, D. 2014: Morphological and Genetic
Diversity of the Wood-Boring Xylophaga (Mollusca, Bivalvia): New Species and Records from Deep-Sea
Iberian Canyons. PLOS One. PDF On-line.
SANTOS, S., TAGLIARO, C., BEASLEY, C., SCHNEIDER, H., SAMPALO, I., FILHO, C. & MÜLLER, A.
2005: Taxonomic implications of molecular studies on Northern Brazilian Teredinidae specimens. Genetics and
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SARASWATHY, M. & NAIR, N. B. 1974: The influence of salinity on a tropical estuarine shipworm Nausitora
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SCHELTEMA, R. S. & TRUITT, R. V. 1956: The shipworm Teredo navalis in Maryland coastal waters.
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SHIPWAY, J. R., BORGES, L. M. S., MÜLLER, J. & CRAGG, S. M. 2014. The broadcast spawning Caribbean
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Shipworm Bankia setacea (Bivalvia: Teredinidae). Applied and Environmental Microbiology. 66(4): 1685-1691.
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Research. 56: 87-94.
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larvae. Wood Research. 59/60: 33-39.
VIKTORAS, D. 2007: Invasive Alien Species Fact Sheet Teredo navalis. On-line database of the North
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WATERBURY, J. B., CALLOWAY, D. B. & TURNER, R. D. 1983: A cellulolytic nitrogen-fixing bacterium
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WOLFF, W. J. 2005: Non-indigenous marine and estuarine species in the Netherlands. Zoologische Mededelingen. 79: 1-116.
8. Family Xylophagaidae HAGA & KASE, 2013
8.1. Systematics
This mono-generic Family is based on its single extant representative:
Genus Xylophaga TURTON, 1822.
Genus Xylophaga TURTON, 1822
Systematics
According to ROMANO, VOIGHT, PEREZ-PORTELA & MARTIN, 2014,
this Genus regroups twelve extant species:
X. dorsalis (TURTON, 1819), X. tipperi TURNER, 2002, X. multichela KNUDSEN, 1961,
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X. depalmai KNUDSEN, 1961, X. guineensis KNUDSEN, 1961, X. mexicana DALL, 1908 (sensu TURNER, 2002),
X. tipperi TURNER, 2002, X. bayeri TURNER, 2002, X. globosa SOWERBY, 1835,
X. japonica TAKI & HABE, 1950, X. indica SMITH, 1904,
and X. multichela ROMANO, VOIGHT, PEREZ-PORTELA & MARTIN, 2008.
Taxonomical remarks
According to WoRMS 2015, thirty-nine species must be added to this list, considered as to restrictive.
It seems that until 2013, no researchers had realized that, in the Regnum Animalia, two Families have been proposed and accepted with identical names: The Family Xylophagidae (Order Diptera: Class Insecta) and the Family Xylophagidae (Order Myoida: Class Mollusca).
Recent solution
The Family Xylophagidae (Insecta: Diptera) having priority by anteriority, the name of the molluscan Family
was modified, in 2013, into Family Xylophagaidae by Haga and Kase.
Singularity
The diverse extant and extinct representatives of this Genus are in possession of a Teredo-like shell
but do not form a calcareous tube.
8.2. Geographical distribution of the Family Xylophagaidae
All the extant representative taxa of this Family inhabit deep to abyssal waters, environments not compatible
with the environment of all the deposits of the earlier Devonian to the Holocene of Belgium.
8.3. Geological record of the Family Xylophagaidae
None
8.4. Ichnology and Xylophagaidae
As far as known, traces of activities of the Xylophagaidae were never mentioned.
They were surely confused with these of the Teredinidae.
8.5. References concerning the Family Xylophagaidae
ANSELL, A. D. & NAIR, N. B. 1969: The mechanisms of boring in Martesia striata Linné (Bivalvia:
Pholadidae) and Xylophaga dorsalis Turton (Bivalvia: Xylophaginidae). Proceedings of the Royal Society of
London, Series B. 174: 123-133.
BORGES, L. M. S., SIVRIKAYA, H., le ROUX, A., SHIPWAY, J. R., CRAGG, S. M. & COSTA, F. O. 2012:
Investigating the taxonomy and systematics of marine wood borers (Bivalvia: Teredinidae) combining evidence
from morphology, DNA barcodes and nuclear locus sequences. Invertebrate Systematics. 26: 572-582
CULLINEY, J. L. & TURNER, R. D. 1976: Larval development of the deep-water wood-boring bivalve,
Xylophaga atlantica Richards (Mollusca, Bivalvia, Pholadidae). Ophelia. 15: 149-161.
DISTEL, D. L. & ROBERTS, S. J. 1997: Bacterial endosymbionts in the gills of the deep-sea wood-boring
bivalves Xylophaga atlantica and Xylophaga washingtona. Biological Bulletin. 192: 253-261.
DONS, C. 1929: Zoologiske Notiser V. Xylophaga dorsalis I Norge. Det Kongelige Norsske Videnskabers
Selskab Forhandlinger. 65: 196-199.
GIRIBET, G. & DISTEL, D. 2003: Bivalve Phylogeny and Molecular data. In: LYDEARD, C. & LINDDBERG,
D., R. Editors. Molecular systematics and phylogeography of mollusks. Washington, D., C. Pp.: 45-90.
HAGA, T. & KASE, T. 2008: Redescription of the Deep-sea wood borer Neoxylophaga teramachii Taki &
Habe, 1950 and its Assignment to the Genus Xyloredo (Bivalvia: Myoida: Pholadoidea) with Comments on
Fossil Pholadoidae. Veliger. 50: 107-119.
HAGA, T. & KASE, T. 2013: Progenetic dwarf males in the deep-sea wood-boring genus Xylophaga (Bivalvia:
Pholadoidea). Journal of Molluscan Studies. 79: 90-94.
HARVEY, R. 1996: Deep water Xylophagaidae (Pelecypoda: Pholadacea) from the North Atlantic with
descriptions of three new species. Journal of Conchology. 35: 473-481.
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JONES, W. J., JOHNSON, S. B., ROUSE, G. W. & VRIJHENHOEK, R. C. 2008: Marine worms (genus
Osedax) colonize cow bones. Proceedings of the Royal Society B. Biological Sciences. 275: 387-391.
KNUDSEN, J. 1961: The bathyal and abyssal Xylophaga (Pholadidae, Bivalvia). Galathea Report. 5: 163-209.
KUDINOVA-PASTERNAK, R. K. 1975: Xylophaga mollusks (Bivalvia, Pholadidae) found in a Scotia Sea
shipwreck. Institute of Oceanology Research Publication. 103: 179-180.
MONARI, S. 2009: Phylogeny and biogeography of pholadid bivalve Barnea (Anchomasa) with considerations
on the phylogeny of Pholadoidea. Acta Palaeontologica Polonica. 54: 315-335.
PURCHON, R. D. 1941: On the biology and relationships of the lamellibranch Xylophaga dorsalis (Turton).
Journal of the Marine Biological Association of the United Kingdom. 25: 1-39.
ROMANO, C., VOIGHT, J. R., COMPANY, J. B., PLYUSCHEVA, M. & MARTIN, D. 2013: Submarine
canyons as a habitat preferred for wood-boring species Xylophaga (Mollusca: Bivalvia). Progress in
Oceanography. 118: 175-187.
ROMANO, C., VOIGHT, J. R., PEREZ-PORTELA, R. & MARTIN, D. 2014: Morphological and genetic
diversity of the wood-boring Xylophaga (Mollusca, Bivalvia): new species and records from deep-sea Iberian
canyons. PLoS One, 9: e102887.
SANTHAKUMARAN, L. N. 1980: Two new species of Xylophaga from Trondheimsfjorden, western Norway.
Sarsia. 65: 269-272.
SANTHAKUMARAN, L. N. & SNELI, J. A. 1984: Studies on the marine fouling and wood-boring organisms
of the Trondheimsfjord (western Norway). Gunneria. 47: 1-36.
SANTOS, S. M. L., TAGLIARO, C. H., BELEYAS, C. R., SCHNEIDER, H., SAMPAIO, I., FILHO, C. S. & de
PAULA MÜLLER A. C. 2005: Taxonomic implications of molecular studies on northern Brazilian Teredinidae
(Mollusca, Bivalvia) specimens. Genetics and Molecular Biology. 28: 175-179.
SHIPWAY, R. J., BORGES, L. S., MÜLLER, J. & CRAGG, S. 2014: The broadcast spawning Caribbean shipworm, Teredothyra dominicensis (Bivalvia, Teredinidae), has invaded and become established in the eastern
Mediterranean Sea. Biological Invasions. 2014: 1-12.
THOMPSON,W. 1847: Note on the Teredo norvegica (T. navalis, Turton, not Linn.), Xylophaga dorsalis,
Limnoria terebrans and Cheluda terebrans, combined in destroying the submerged wood-work at the harbor of
Ardrossan on the coast of Ayrshire. Annals and Magazine of Natural History. 20: 157-164.
TURNER, R. D. & CULLINEY, J. 1971: Some anatomical and life history studies of wood boring bivalve systematics. Annual Report of the American Malacological Union for 1970.1971: 65-66.
TURNER, R. D. 1955: The fanily Pholadidae in the western Atlantic and the eastern Pacific. Part II – Martesesiinae, Jouannetiinae and Xylopagainae. Johnsonia. 3: 65-160.
TURNER, R. D. 2002: On the subfamily Xylophagainae (family Pholadidae, Bivalvia, Mollusca). Bulletin of
the Museum of Comparative Zoology. 157: 223-307.
VOIGHT, J. R. 2007: Experimental deep-sea deployments reveal diverse Northeast Pacific wood-boring
bivalves of Xylophagainae (Myoida: Pholadidae). Journal of Molluscan Studies. 73: 377-391
VOIGHT, J. R. 2008: Deep-sea wood-boring bivalves of Xylophaga (Myoida: Pholadidae) on the Continental
Shelf: a new species described. Journal of the Marine Biological Associon of the United Kingdom. 88: 14591464.
VOIGHT, J. R. 2009: Near-shore and offshore wood-boring bivalves (Myoida: Pholadidae: Xylophagainae) of
the deep Eastern Pacific Ocean: diversity and reproduction. Journal of Molluscan Studies. 75: 167-174.
VOIGHT, J. R. & SEGONZAC, M. 2012: At the bottom of the deep blue sea: a new wood-boring bivalve
(Mollusca, Pholadidae, Xylophaga) from the Cape Verde Abyssal Plain (subtropical Atlantic). Zoosystema. 34:
171-180.
9. Data furnished by the wood pieces
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The data furnished by the examination of the fossil teredinid tubes discovered in the Sint-Niklaas Phosphorite
Bed are summarised in the following paragraphs.
9.1. Nature of these pieces of wood
Except for the fossil illustrated on Plate 8*, all the other pieces of wood are parts of Gymnospermae and,
according to their wood structure, attributable to fossil representatives of the Order Pinales G OROZHANKIN, 1904.
*The piece of wood illustrated on this plate could be a fragment of an adventive root of a mangrove vegetal taxon.
The presence of the Order Pinales in this level of Lower Oligocene Age is confirmed by a detailed microscopic
examination of one of these pieces of wood *, realised by Hugues Doutrelepont**, which will be published in the
volume devoted to the study of the vegetal remains discovered in this level (Géominpal Belgica. 5.4.).
*Precision: Genus Pinus, Family Pineaceae. **M.R.A.C., Tervuren, Belgium.
Pinaceae are common trees on diverse continents, but also, independently of the climate, along a large variety of
coastal zones.
9.2. Degradation of these pieces of wood
These pieces of wood were originally parts of living trees. All these pieces of wood seem to have been parts of
healthy trees. Only some of these present traces of activities of larvae of xylophagous Insecta *.
*These traces will be described in: Addition 3 to Géominpal Belgica. 5.2.
The scarcity of these traces, as well as the scarcity of fungal activities, allows suggesting that their death cannot
be attributed to these continental living forms.
Additionally, none of these pieces of wood present traces of fire, fact that allows considering that their death
cannot be attributed to forest fires.
The senior-author would not be surprised to discover that the event having caused their death, was simply a
tsunami. Tsunamis are responsible for important land destructions, but also for the destruction of large areas of
sandy coasts.
During the Oligocene Period, the area, now named Belgium, is supposed only having had clayish and sandy
coasts in a radius of forty kilometres of Sint-Niklaas.
9.3. Duration of their drifting
Pieces of wood may drift for days to more than tens of years. The longer the duration of their drifting is, the
longer the time is the teredinid bivalves dispose of to grow in these pieces of wood. But the average of growing
varies for each specific taxon: as far as known, from some millimetres per year to three centimetres per year.
9.4. First embedding
The pieces of wood carrying their teredinid hosts were dis-embedded from a first sedimentation place and on a
beach where they were quickly covered by fine sediments. The quality of preservation of their tubes and this of
the wood pieces are the arguments that allow supposing that this supposition is right.
9.5. Power of the dis-embedding flow
Once petrified, these fossil wood-pieces are relatively heavy fossils*. But, if originally, embedded in silty-sandy
deposits, the abrasive effect of a slow water stream may be sufficient to extract these fossils.
*Only the siderite concretions enveloping the undetermined species of the Family Cerianthidae M ILNE-EDWARDS &
HAIME, 1852 are heavier masses.
9.6. Filling of the teredinid tubes
More or less shortly* after the wood pieces stranded, these pieces of wood dried and their hosts putrefied or
mummified**.
*The hotter and drier the surface temperature of the beach was, the more quickly they dried.
**In the thin pieces of wood, a mummification was possible.
The absence of small invertebrate* remains in these fossil tubes signifies that these ones did not have time en-
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ough to reach this source of food, because the sedimentological enveloping was very rapid.
Such as the cleaning of some tubes demonstrated, they were filled by fine sand cores which, principally,
agglomerated around the anterior part of the body of their Teredo-like builders.
9.7. Second embedding
All these teredinid tubes were discovered, in some petrified pieces of wood, in the southern part of the Clay Pit
and extracted from the lower part of the Sint-Niklaas Phosphorite Bed.
The senior-author is still searching a plausible explanation for the absence of drifted pieces of petrified wood in
the upper part of the Sint-Niklaas Phosphorite Bed.
The Sint-Niklaas Phosphorite Bed is a discontinued level constituted by large and irregular lenses of siderite
concretions, which was formed shortly before the formation of the huge banks of the oysters Pycnodonte
callifera de LAMARCK, 1819.
It seems that the environment* was too enclosed to allow the entrance of animals larger than decapod
crustaceans, some gastropod mollusks and some small teleostean fishes.
*Top of large siderite concretions and large sandy places covered by marine to brackish waters during the tidal hours.
9.8. Geochemical alteration of the filling of the tubes
Diverse fossils* contain, in variable quantity, iron-sulfides which may completely destroy them by oxidation.
The filling material of the teredinid tubes may also be affected by this process, which explains the dark colour of
their cross sections.
*All the siderite concretions enveloping the emerging part of the cerianthid tubes, all the siderite concretions enveloping the
burrows of diverse stomatopod crustaceans and all the vertebrate remains.
This geochemical alteration of the sediment filling their tubes was an oxidation phase. This phenomenon is only
possible when such specimens are out of the water, which means that it happened during the stranding phase or
just after the filling phase.
Just after the oxidization* of the sediment filling the teredinid tubes, these pieces of wood were covered by an
accumulation of muddy and sandy sediments mixed with other heavy siderite concretions.
*Partial: Some tubes contain unaffected cores.
10. Conclusions
10.1. Systematic conclusion
The attempt to precise the systematic attribution of these rare fossils is, of course, an interesting aspect, but the
existence of such fossils in older and younger strata being a certitude, their discovery is a relatively poor addition
to the Natural History of these ultra-specialised bivalves.
Their tubes are made of calcium carbonates. They are approximately rectilinear *, never scrolled or regularly
curved.
*Except when approaching a wood-node.
The cross-section of these tubes is sub-circular. Their diameter varies from four to nine millimetres. The
maximal length observed is 23,4 centimetres.
The very restricted distribution zones of some extant generic taxa could allow excluding the possibility that these
ones are related to the Belgian Lower Oligocene teredinids discovered in the Sint-Niklaas Phosphorite Bed.
For example: The extant Genus Bankia is represented by two species endemic to the eastern Pacific Ocean.
But during the Lower Oligocene*, a passage across Central America was possible for any hosted drifting pieces
of wood as well as for larvae of any teredinid-species.
*And this crossing remains possible till to the earlier Pliocene.
Only one tube seemed * in possession of the pallets of its builder. The morphology of these pallets is very similar
to this of the extant representatives of the Genus Bactrophorus TAPPARONE CANEFRY, 1877, but considering the
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variation of the morphology of their pallets during the growth of their owner, the senior-author considers that an
attribution to this Genus will also be doubtful.
*In fact, these fossil remains were the cross-sections of the two valves of a little Bivalvia, having inhabited an empty tube.
The extant Genera including species that inhabit fresh- or brackish- waters cannot be excluded because nobody
knows when their first fresh- or brackish- members appeared.
Only the deep to abyssal distribution of the extant representatives of the Family Xylophagaidae seems allowing
to suppose they have no phylogenetic relations with the Belgian Lower Oligocene teredinids discovered in the
Sint-Niklaas Phosphorite Bed.
With so few data, the only honnest systematic approach seems to be: Family Teredinidae, Genus indet, sp. indet.
The senior-author considers that the list of invalid teredinid taxa is sufficiently long to avoid adding one more
taxon with the unique argument or diagnosis: It is a new teredinid taxon, thirty-two million years aged, of which
the tubes may present obvious transversal rings.
But, it seems possible to precise, a little more, the systematic position of these fossils (See further: Chapter 13).
And the systematic position of two Families Teredinidae and Xylophagaidae, such as this of their nearest
parents, need to be reconsidered. See further: Chapter 13).
10.2. Taphonomical interpretation
The fact that none of these tubes are flattened allows suggesting that their petrification preceeded their first
embedding.
Their examination demonstrates that they were successively petrified, embedded, dis-embedded and regrouped
in the lower part of some lenses of the Sint-Niklaas Phosphorite Bed.
Some of these pieces of wood also present different traces of activities of larvae of xylophagous insects. The
sole period during which these larvae had the possibility to feed upon diverse parts of these trees was when growing on emerged surfaces.
11. Special adaptations of some Bivalvia
11.1. General considerations
The following morphological considerations may seem more than obvious but the senior-author points out these
ones, because once genetically encoded, such significant morphological modifications are irreversible.
After examination of some of these significant morphological modifications and their ecological implications, it
will be interesting to know when* they appeared and which could be the extern causes ** of their apparition.
*Geologically speaking.
**Astrophysical causes.
11.2. Identic or different valves
The majority of the extant and extinct, taxa of the Bivalvia possess two perfectly symmetric valves. But other
ones, such as all the representatives of the Families * Ostreidae, Anomiidae and Pectinidae, possess two valves of
different size and morphology and live in icy waters to equatorial waters from the intertidal zone to abyssal
zones. *List non exhaustive.
The fact that these Families are represented by many taxa in all parts of the world and at various depths allows
supposing that the climate was never responsible for their morphological changes nor for their genetic code
modifications.
11.3. Advantages presented by the possession of identic valves
A symmetric morphology of the valves has for principal, if not unique, advantage the possibility to move regularly in the same direction in all types of sediments or bored supports.
If stranded by storm turbulences, many of these molluscs seem to be able to escape from deshydration by a very
rapid return in sediments covered by waters.
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11.4. Advantages presented by the possession of different valves
The massive and deep ventral valve of the free-living* taxa of the Family Ostreidae allows a partial embedding
in the sediment surrounding their shell and ensure a relative stability on the sea bottom.
*Many other taxa of this Family live, or lived, strongly fixed on diverse inorganic or organic supports .
The aerodynamic morphology of the valves of the extant and extinct Pectinidae allows them to realize a retropropulsion comparable to this of many cephalopods.
The morphology of the shell of all the representative taxa of the Family Anomiidae allows them to realize
different very efficient types of fixation on irregular supports.
11.5. Disadvantages presented by the by the possession of different valves
If the turbulence of the water increases too much, the position of the shell of the free-living Ostreidae becomes
unstable and can be inversed, which ineluctably induces the death of the animal. The formation of large banks
allows reducing this risk, but never completely.
The turbulence of the water has very few implications for the Pectinidae, but during their retro-saltation, the very
large angle of their visibility remains insufficient to avoid some obstacles or other large predators positioned in
their dead angle of visibility.
The Anomyidae are sessile molluscs and depend essentially of the stability or the longevity of their support.
12. Short examination of the principal data concerning the diverse Families
phylogenetically considered as parents of the Teredinidae
12.1. Motivation
A short examination of the principal data concerning the diverse Families classically included in the
Pholadomyoidea, Myoidea, Hyatelloidea, Thracioidea and Clavagelloidea allows suggesting that their systematic is not coherent.
The systematic of the following Families: F. Myidae de LAMARCK, 1809, F. Myochamidae CARPENTER, 1860, F.
Corbulidae de LAMARCK, 1818, F. Pandoridae RAFINESQUE, 1815, F. Laternulidae KING & BRODERIP, 1832, F.
Lyonsiidae FISCHER, 1887, F. Periplomatidae DALL, 1895, F.Thraciidae STOLICZKA, 1870, F. Verticordiidae
STOLICZKA, 1870, F. Cuspidariidae DALL, 1886, F. Teredinidae RAFINESQUE, 1815, F. Xylophagaidae HAGA &
KASE, 2013, F. Hiatellidae GRAY, 1824, F. Pholadidae de LAMARCK, 1809, F. Gastrochaenidae GRAY, 1840, F.
Clavagellidae d’ORBIGNY, 1843, F. Penicillidae BRUGUIERE, 1789 and F. Pholadomyiidae KING, 1844, is briefly
reviewed and commented in the following paragraphs.
12.2. Recent revision
In 2010, Bieler, Carter and Coan * proposed a new classification for the Bivalvia and suggested Pholadomyida
NEWELL, 1965 in place of Anomalodesmata ** DALL, 1889, but the responsibles of WoRMS have choised the
second option.
*BIELER, R., CARTER, J. G. & COAN, E. V. 2010: Classification of Bivalve families. Pp.: 113-133 In BOUCHET, P. &
ROCROI, J.-P. 2010: Nomenclator of Bivalve Families. Malacologia. 52(2): 1-184.
**The first Anomalodesmata appeared during the Middle Ordovician Period, circa 560 million years ago.
WoRMS 2015
also proposed to regroup into
eigth super-Families all the extant representatives of the Anomalodesmata:
1.The super-Family Clavagelloidea, reprouping the two Families:
Clavagellidae and Penicillidae.
2.The super-Family Cuspidarioidea, reprouping the two Families:
Cuspidariidae and Spheniopsidae.
3. The super-Family Myochamoidea, reprouping the two Families:
Myochamidae and Cleidothaeridae.
4. The super-Family Pandoroidea, reprouping the two Families:
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Pandoridae and Lyonsiidae.
5. The super-Family Pholadomyoidea, based on the Family:
Pholadomyidae.
6. The super-Family Poromyoidea, based on the Family:
Poromyiidae.
7. The super-Family Thracioidea, regrouping the three Families:
Thraciidae, Laternulidae and Periplomatidae.
8. The super-Family Verticordioidea, regrouping the two Families:
Verticordidae and Lyonsiellidae.
12.3. Remarks
Some malacologists add the Family Halonymphydae* SCARLATO & STAROBOGATOV, 1983 and the Family
Protocuspidariidae** SCARLATO & STAROBOGATOV, 1983 in the super-Family Cuspidarioidea and the Family
Euciroidae*** DALL, 1895 in the super-FamilyVerticordioidea.
*The Family Halonymphydae SCARLATO & STAROBOGATOV, 1983 for the Genus Halonympha DALL & SMITH, 1886.
**The Family Protocuspidariidae SCARLATO & STAROBOGATOV, 1983 for regrouping the two Genera: Protocuspidaria
ALLEN & MORGAN, 1981 and Multitentacula KRYLOVA, 1995.
***The Family Euciroidae DALL, 1895 is proposed for regrouping the two Genera: Genus Euciroa DALL, 1881 and Genus Acreuciroa THIELE & JAECKEL, 1931.
12.4. Family Myidae de LAMARCK, 1809
Systematics
According to WoRMS 20015, this Family regroups the six extant Genera:
Genus Mya LINNAEUS, 1758, Genus Sphenia TURTON, 1822, Genus Platyodon CONRAD, 1837,
Genus Tugonia GRAY, 1842, Genus Cryptomia CONRAD, 1848 and Genus Paramya CONRAD, 1860.
Remark
The validity of the Genus Tugonella JOUSSEAUME, 1891 remains subject of controversy.
Morphological and environmental adaptations
All the extant representatives of this Family possess symmetric valves and one pair of, more or less elongated,
siphons. As far as known their extinct representatives also lived buried in the sediment.
For the representative specific taxa of these Genera, the simplest and the best solution for escaping to extern
physical aggressions seems to have been to live deeper and deeper in the sediment: The increasing of the mass of
the sediments in which they live has for result the decrease of the possibility to enter into contact with all the
particles able to damage their genetic code.
An increasing of the size of their siphon and an enforcement of the power of their foot are sufficient to realize
this performance.
In Belgium, Paleoichnology demonstrates that since the Upper Eocene, some populations of extinct representtatives of this Family bored deeper and deeper * galleries.
*Particularly in sandy sediments, because muddy sediments always offer a better protection against ionised or radioactive
particles.
Some references for Family Myidae
BIELER, R., CARTER, J. G. & COAN, E. V. 2010: Classification of Bivalve families. See: 113-133. In: BOUCHET, P. & ROCROI, J.-P. 2010: Nomenclator of Bivalve Families. Malacologia. 52(2): 1-184.
COAN, E. V. 1999: The eastern Pacific species of Sphenia (Bivalvia: Myidae). The Nautilus. 113(4): 103-120.
Integrated Taxonomic Information System 2013: Myidae.
LAMPRELL, K. & STANISIC, J. 1998: Myidae from Australian waters (Mollusca, Bivalvia). Molluscan
Research. 19(1): 1-10.
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MORRIS, R. H., ABOTT, D. P. & HADERLIE, E. C. 1980: Intertidal Invertebrates of California. XV + 690 p.,
200 pls. Stanford University Press.
OLIVIER, P. G. & CHESNEY, H. C. G. 1997: Taxonomy and descriptions of bivalves (Lucinoidea,
Galeommatoidea, Carditoidea, Cardioidea, Tellinoidea & Myoidea) from the Arabian Sea. Journal of
Conchology. 36(1): 51-76.
PASTORINO, G. & BAGUR, M. 2011: The genus Sphenia Turton, 1822 (Bivalvia: Myidae) from shallow
waters of Argentina. Malacologia. 84(1-2): 431-135.
ZHANG, J. L., XU, F. S. & LIU, J. Y. 2012: The Myidae (Mollusca, Bivalvia) from Chinese waters with
description of a new species. Zootaxa. 3383: 39-60.
12.5. Family Myochamidae CARPENTER, 1861
(Plate 24)
Systematics
According to the researches of the senior-author, this Family includes the four Genera:
Genus Myochama STUTCHBURY, 1830, Genus Myadora GRAY, 1840,
Genus Hunkydora FLEMING, 1948 and Genus Myadoropsis HABE, 1960.
Ecology and environment
According to WoRMS 2015, this Family only regroups taxa inhabiting marine waters.
All their representatives were discovered in sandy to silty bottoms,
from the coastal zone to the upper part of the continental shelf.
As far as known, all the extant representatives of this Family were discovered in the eastern Pacific,
from Japanese waters to Australian and New Zealand waters.
Geological record
Upper Eocene to Recent Times.
Some references for Family Myochamidae
BEU, A. G. 2006: Marine Mollusca of oxygen isotope stages of the last 2 million years in New Zealand. Part 2.
Biostratigraphically useful and new Pliocene to recent bivalves. Journal of the Royal Society of New Zealand.
36(4): 151-338. See p.: 311.
FLEMING, C. A. 1948: New species and genera of marine Mollusca from the Southland Fiords. Transactions of
the Royal Society of New Zealand. 77: 72-92. See p.: 80.
HABE, T. 1960: New species of molluscs from the Amakusa Marine Biological Laboratory, Reihoku-cho, Amakusa, Kumamoto Prefecture, Japan. Publications of the Seto Marine Biological Laboratory. 8(2): 289-298.
HUBER, M. 2010: Compendium of bivalves. A full-color guide to 3,300 of the world’s marine bivalves. A status on Bivalvia after 250 years of research. Hackenheim: ConchBooks. 901 pp.
MARSHALL, B. A. 2002: Some Recent Thraciidae, Periplomatidae, Myochamidae, Cuspidariidae and
Spheniopsidae (Anomalodesmata) from the New Zealand region and referral of Thraciareinga Crozier, 1966 and
Scintillona benthicola Dell, 1956 to Tellimya Brown, 1827(Montacutidae) (Mollusca: Bivalvia). Molluscan
Research. 22(3): 221-288.
VERMEIJ, G. J. 2005: From Europe to America: Pliocene to Recent trans-Atlantic expansion of cold-water
North-Atlantic molluscs. Proceedings of the Royal Society. Biological Sciences. 272(1580): 2545-2550.
STUTCHBURY, S. 1830: On two new genera of testaceous mollusca,and five new species of the genus Anatina,
lately discovered at Port Jackson, New South Wales. Zoological Journal. 5: 95-101, pl.: 42.
12.6. Family Corbulidae de LAMARCK, 1818
Systematics
According to the researches of the senior-author, this Family could regroup the thirteen extant Genera:
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Genus Corbula BRUGUIERE, 1797, Genus Lentidium de CRISTOFORI & JAN, 1832,
Genus Anticorbula DALL, 1898, Genus Caestocorbula VINCENT, 1910, Genus Caryocorbula GARDNER, 1926,
Genus Anisocorbula IREDALE, 1930, Genus Varicorbula GRANT & GALE, 1931,
Genus Hexacorbula OLSSON, 1932, Genus Tenuicorbula OLSSON, 1932, Genus Panamicorbula PILSBRY, 1932,
Genus Juliacorbula OLSSON & HARBISON, 1953, Genus Potamocorbula HABE, 1955 and
Genus Apachecorbula OLIVEIRA, 2014.
Systematic remarks
The Genus Apachecorbula* OLIVERA, 2014 is not mentioned by WoRMS 2015, and
the validity of the Genus Potamocorbula HABE, 1955 remains controversial.
Reflexion of a paleontologist
The extreme differences of the morphology * of the shell of the diverse extant species
attributed to the Genus Corbula makes difficult to understand its malacological signification.
*See Genus Corbula on the site Shell Encyclopedia.
For the representative specific taxa of these Genera, the construction of a shell presenting two valves of different
sizes, the left one being obviously of smaller size than the right one, allowed them to close strongly their shell,
which considerably increased the protection of their body.
The populations of the nine specific taxa attributed to the Genus Potamocorbula HABE, 1955 inhabit brackish
waters. E.g.: C. amurensis (SCHRENCK, 1861) is a species inhabiting marine and brackish waters in the northern
Pacific Ocean. Its distribution includes Siberia to China, Korea and Japan.
In Belgium, it is only after the Eocene-Oligocene Transition, that diverse representative taxa of the Genus
Corbula sensu BRUGUIERE, 1797 appeared.
Some references for Family Corbulidae
BOUCHET, P. & ROCROI, J.-P. 2010 : Nomenclator of bivalve families; with a classification of bivalve families by BIELER, R., CARTER, J. G. & COAN, E. V. Malacologia. 52(2): 1-184.
mollusks from Baja California to northern Peru. 2 vols. 1258 p.
KORNIUSHIN, A. V. & GLAUBRECHT, M. 2003: Novel reproductive modes in freshwater clams: brooding
and larval morphology in Southeast Asian taxa of Corbicula (Mollusca, Bivalvia, Corbiculidae). Acta Zoologica.
84(4): 293-315.
Integrated Taxonomic Information System 2014: Corbulidae.
SHEPPARD, A. 1984: The molluscan fauna of Chagos (Indian Ocean) and an analysis ot its broad distribution
patterns. Coral Reefs. 3: 43-50.
12.7. Family Pandoridae RAFINESQUE, 1815
(Plate 29)
Systematics
According to WoRMS 2015, this Family regroups the six extant Genera:
Genus Pandora BRUGUIERE, 1797, Genus Cliodiophora CARPENTER, 1864, Genus Heteroclidus DALL, 1903,
Genus Foveadens DALL, 1915, Genus Frenamya IREDALE, 1930 and
Genus Coania VALENTICH-SCOTT & SKOGLUND, 2010.
Systematic remark
The huge morphological variability* of the shell of the specific taxa attributed to these Genera,
such as this of numerous Genera of the Myoidea makes perplex any paleomalacologists.
*E.g.: See Genus Pandora or Genus Corbula in the Shell Encyclopedia, published by Guido and Philippe Poppe.
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Taxonomic remark
The Genus Pandora is also a Genus of Fungi, Order Entomophthorales (Zygomycaeta).
Another constation which illustrates the lack of contacts between Biologists.
Generalities
The Family Pandoridae is represented by species in possession of a strongly compressed shell. Their populations
inhabit mostly cold marine waters from the littoral zone to areas of more than five hundred metres depth. These
lasts* have a very thin and translucid shell.
*Such as Pandora pinna (MONTAGU, 1803).
This Family is world-widely distributed.
Geological records
The first fossils attributable to this Family were discovered in strata of Ypresian Age (Lower Eocene).
Some references for Family Pandoridae
BOSS, K. J. & MERRILL, A. S. 1965: The family Pandoridae in the western Atlantic. Johnsonia. 4:181-215,
pls. 115-126.
CARPENTER, P. P. 1865: Contributions towards a monograph of the Pandoridae. Proceedings of the Zoological
Society of London. 1864: 596-603.
DOGAN, A., ÖNEN, M., ÖZTÜRK, B & BITLIS, B. 2008: On the presence of Pandora inaequivalvis
(Bivalvia: Pandoridae) on the Levantine coast of Turkey. Marine Biodiversity Records. 1: Electronic publication.
MORTON, B. 1984: The adaptations of Frenamya ceylanica (Bivalvia: Anomalodesmata: Pandoracea) to life on
the surface of soft muds. Journal of Conchology. 31: 359-371.
THOMAS, K. A. 1994: The functional morphology and biology of Pandora filosa (Carpenter, 1864) (Bivalvia:
Anomalodesmata: Pandoracea). The Veliger. 37: 23-29.
VALENTICH-SCOTT, P. & SKOGLUND, C. 2010: A review of the Recent Pandoridae (Bivalvia) in the
Panamic Province, with descriptions of three new species. The Nautilus. 124(2): 55-76.
VERMEIJ, G. J. 1987: Evolution and Escalation: An Ecological History of Life. Princeton University Press,
Princeton, New Jersey. 527 p.
YONGE, C. M. & MORTON, B. 1980: Ligament and lithodesma in the Pandoracea and the Poromyacea with a
discussion on evolutionary history in the Anomalodesmata (Mollusca: Bivalvia). Journal of Zoology. 191: 263292.
12.8. Family Laternulidae KING, 1832
Systematics
According to the researches of the senior-author, this Family is based on its generotype:
Genus Laternula RÖDING, 1798 and only includes a second Genus: Genus Clistoconcha SMITH, 1910.
Generalities
The type-species of the Genus Laternula, L. elliptica (KING & BRODERIP, 1832) is a filter feeder. Its individuals
inhabit lightly embedded in the sea bottom with their siphons extended to the surface.
The diverse populations of Laternula elliptica known live in the Southern Ocean, around Antarctica and the tip
of Patagonia. It is usually found in shallow waters but the greatest depth at which it has been recorded is 360
metres.
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They live in soft bottoms, constituted by muddy sand and gravel, in which they deeply embed. Their
concentrations rise frequently up to a hundred individuals per square metre.
Geological records
Pliocene fossils of Laternula elliptica are very common in some sedimentary rocks on the Cockburn Island and
the James Rock Island off the Antartic Peninsula.
Some fossils of Rhaetian Age are considered as their oldest ancestors.
Some references for Family Laternulidae
HUBER, M. 2010: Compendium of bivalves. A full-color guide to 3.300 of the world’s marine bivalves. A status on Bivalvia after 250 years of research. Hackenheim: ConchBooks. 901 p.
KING, P. P. 1832: Description of the Cirrhipeda, Conchifera and Mollusca, in a collection formed by the officers
of H.M.S. Adventure and Beagle employed between the years 1826 and 1830 in surveying the southern coasts of
South America, including the Straits of Magalhaens and the coast of Tierra del Fuego. Zoological Journal.5: 332
to 349.
MELVILL, J. C. & STANDEN, R. 1914: Notes on Mollusca collected in the north-west Falklands by Mr.
Rupert Valentine, F. L. S., with descriptions of six new species. Annals and Magazine of Natural History. 8(13):
110-136, pl. 7.
PETIT, R. E. 2009: George Brettingham Sowerby, I, II & III: their conchological publications and molluscan
taxa. Zootaxa. 2189: 1-218.
12.9. Family Lyonsiidae FISCHER, 1887
Systematics
According to Gofas (2010), this Family regroups the four Genera:
Genus Lyonsia TURTON, 1822, Genus Entodesma PHILLIPS, 1845,
Genus Allogramma DALL, 1903 and Genus Bentholyonsia HABE, 1952.
According to WoRMS 2015, this Family regroups the four Genera:
Genus Lyonsia TURTON, 1822, Genus Entodesma PHILIPPI, 1845, Genus Mytilimeria CONRAD, 1837 and,
with reserves, Genus Sinolyonsia* XU, 1992.
All the fourtheen other Genera, proposed between 1822 and 1903, are considered as synonyms
of the Genera Allogramma, Entodesma or Lyonsia.
Singular taxonomic unprecisions
According to WoRMS 2015, within Family Lionsyiidae, the Genus Hiatella is considered as a synonym of
Genus Lyonsia, but when searching, on WoRMS-site, for Genus Hiatella,
this Genus is considered as valid and regroups four extant species.
Generalities
All the extant representative taxa of this Family have inequivalve and oblong valves. Their right valve is more
convex than their left valve.The ligament connecting their two valves is located in an internal groove.
The Family is world-widely distributed, but some of its specific taxa
are endemic to, more or less, large areas:
E.g.:
Allogramma formosa (JEFFREYS, 1882), Entodesma brasiliense (GOULD, 1850),
Entodesma patagonicum (d'ORBIGNY, 1846), Lyonsia californica CONRAD, 1837,
Lyonsia floridana CONRAD, 1849, Lyonsia malvinensis d'ORBIGNY, 1846,
Lyonsia norwegica (GMELIN, 1791), Lyonsia panamensis DALL, 1908
or Lyonsia taiwanica LAN & OKUTANI, 2002.
Geological records
Thanetian Age (Paleocene) to Present Times.
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Some references for Family Lyonsiidae
GOFAS, S., LE RENARD, J. & BOUCHET, P. 2001: Mollusca. In: Costello, M. J., EMBLOW, C. & WHITE,
R. J. Eds.: 2001: European register of marine species: a check-list of the marine species in Europe and a
bibliography of guides to their identification. Collection Patrimoines Naturels. 50: pp. 180-213.
LAN, T. C. & OKUTANI, T. 2002: A new bivalve Lyonsia taiwanica from mangrove swamp near Tainan,
Taiwan. Memoirs of the Malacological Society of Taiwan. 3: 26-29.
MACSOTAY, O. & CAMPOS, R. Eds. 2001: Moluscos representativos de la plataforma de Margarita,
Venezuela. Valencia, Venezuela, III + 280 p., 32 pls.
PIMENTA, A. D. & OLIVEIRA, C. D. 2013: Taxonomic Review of the Genus Lyonsia (Pelecypoda:
Lyonsiidae) from East Coast of South America, with Description of a New Species and Notes on Other Western
Atlantic Species. American Malacological Bulletin. 31(1): 75-84.
12.10. Family Periplomatidae DALL, 1895
Systematics
According toWoRMS 2015, this Family is based on the extant taxon:
Genus Periploma SCHUMACHER, 1817 and includes the five other extant generic taxa:
Genus Cochlodesma COUTHOUY, 1839, Genus Halistrepta DALL, 1904,
Genus Offadesma IREDALE, 1930, Genus Albimanus PILSBRY & OLSSON, 1935 and
Genus Pendaloma IREDALE, 1930.
Genus Periploma SCHUMACHER, 1817
This Genus is supposed to regroup some thirty-one extant specific taxa*:
P. margaritaceum (de LAMARCK, 1801), P. papyratium (SAY, 1822), P. leanum (CONRAD, 1831),
P. lenticulare SOWERBY I, 1834, P. planiscula SOWERBY I, 1834, P. fragile (TOTTEN, 1835),
P. inequale (ADAMS, 1842), P. compressum d’ORBIGNY, 1846, P. ovatum d’ORBIGNY, 1846,
P. orbiculare GUPPY, 1882, P. aleuticum (KRAUSE, 1885), P. discus STEARNS, 1890, P. carpenteri DALL, 1896,
P. stearnsii DALL, 1896, P. indicum MELVILL, 1898, P. andamanicum (SMITH, 1904),
P. teevani HERTLEIN & STRONG, 1946, P. lagartillum OLSSON, 1961, P. fracturum BOSHOFF, 1968,
P. coquettae VAN REGTEREN ALTENA, 1968, P. subfragilis SCARLATO & KAFANOV, 1988,
P. rosewateri BERNARD, 1989, P. camerunense COSEL, 1995, P. sanctamarthaense ARDILLA & DIAZ, 1998,
P. beibuwanse XU, 1999, P. multigranosum XU, 1999, P. nanshaense XU, 1999,
P. hendrickxi VALENTICH-SCOTT & COAN, 2010, P. kaiserae VALENTICH-SCOTT & COAN, 2010 and
P. skoglundae VALENTICH-SCOTT & COAN, 2010.
Or, if referencing to WoRMS 2015, more than seventy specific taxa.
But,the control of the highly controversed validity of these taxa is not the aim of this publication.
Generalities
The extant representatives of this Genus are middle-sized (two to four centimetres length) marine bivalves
presenting a shell without hinge and inhabit shallow marine waters in diverse parts of the world.
The Genus has a very large dispersal, but many of its representatives are endemic to restricted areas, which some
speficic names obviously indicate.
E.g.:
P. aleuticum, P. andamanicum, P. beibuwanse, P. camerunense,
P. indicum, P. nanshaense and P. sanctamarthaense.
Geological records
The specimens of P. macphersoni MARWICK, 1931, discovered in
New-Zealand, seem to be their oldest representatives.
The Genus Offadesma IREDALE, 1930 is represented by some species discovered
in diverse strata dating from the Miocene Age in Patagonia (Argentina).
Some references for Family Periplomatidae
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mollusks from Baja California to northern Peru. 2 vols, 1258 pp.
GRIFFIN M PASTORINO G 2006: The Genus Offadesma Iredale, 1930 (Bivalvia: Periplomatidae) in the
Miocene of Patagonia. The Veliger. 48(2): 75-82 .
MARWICK, J. 1931: The Tertiary Mollusca of the Gisborne district. Journal Paleontological Bulletin of NewZealand. 13: 1-177.
MAXWELL, P. A. 2009: Cenozoic Mollusca. Pp.: 232-254. In: GORDON, D. P. Ed. 2009: New Zealand
inventory of biodiversity. Volume One: Kingdom Animalia: Radiata, Lophotrochozoa, Deuterostomia.
Canterbury University Press, Christchurch.
12.11. Family Thraciidae STOLICZKA, 1870
(Plate 21)
Systematics
According to WoRMS 2015, this Family regroups the eleven extant Genera:
Genus Thracia LEACH in de BLAINVILLE, 1824, Genus Cyathodonta CONRAD, 1849,
Genus Asthenothaerus CARPENTER, 1864, Genus Bushia DALL, 1886, Genus Parvithracia FINLAY, 1926,
Genus Lampeia MACGINITIE, 1959, Genus Trigonothracia YAMAMOTO & HABE, 1959,
Genus Pseudocyathodonta* COHAN, 1990, Genus Skoglundia* COHAN, 1990,
Genus Barythaerus MARSHALL, 2002 and Genus Pelopina HUBER, 2010.
Systematic remark
The validity of the five following extant Genera remains controversed:
Genus Phragmorisma TATE, 1894, Genus Thracidora IREDALE, 1924, Genus Cetothrax IREDALE, 1949,
Genus Thracidentula GARRARD, 1961 and Genus Thraciopsis TATE & MAY, 1900.
Generalities
The extant representatives of this Family are saltwater bivalves inhabiting cold waters.
Their shell possesses no tooth on their hinge and their length varies from three to eigth centimetres.
The populations of its numerous specific taxa are scattered
in the North Est Atlantic Ocean, including the North Sea and the Mediterranean Sea,
in the North West Atlantic Ocean, including the Carribean Sea,
and in the eastern part of the Pacific Ocean (Japan, New Zealand, Australia, Antarctic).
Geological records
Rhetian (Lower Triassic) to Present Times.
Some references for Family Thraciidae
FINLAY, H. J. 1926: A further commentary on New Zealand molluscan systematics. Transactions of the New
Zealand Institute. 57: 320-485.
HUBER, M. 2010: Compendium of bivalves. A full-color guide to 3,300 of the world’s marine bivalves. A
status on Bivalvia after 250 years of research. Hackenheim: ConchBooks. 901 p. See: Thraciidae.
MARSHALL, B. A. 2002: Some Recent Thraciidae, Periplomatidae, Myochamidae, Cuspidariidae and
Spheniopsidae (Anomalodesmata) from the New Zealand region and referral of Thraciareinga Crozier, 1966 and
Scintillona benthicola Dell, 1956 to Tellimya Brown, 1827(Montacutidae) (Mollusca : Bivalvia). Molluscan
Research. 22(3): 221-288.
SARTORI, A. F. & DOMANESCHI, O. 2005: The functional morphology of the Antarctic bivalve Thracia
meridionalis Smith, 1885 (Anomalodesmata: Thraciidae). Journal of Molluscan Studies. 71: 199-210.
12.12. Family Cuspidariidae DALL, 1886
(Plate 21)
Systematics
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According to WoRMS 2015, this Family regroups the twenty following extant Genera:
Genus Cuspidaria NARDO, 1840, Genus Myonera DALL in GRIFFITH & PIDGEON, 1834,
Genus Cardiomya ADAMS, 1864, Genus Leiomya ADAMS, 1864, Genus Plectodon CARPENTER, 1865,
Genus Vulcanomya DALL, 1886, Genus Rhinoclama DALL & SMITH, 1886,
Genus Tropidomya DALL & SMITH, 1886, Genus Luzonia DALL & SMITH in DALL, 1890,
Genus Pseudoneaera STURANY, 1901, Genus Austroneara POWELL, 1937,
Genus Jeffreysomya NORDSIECK, 1969, Genus Rengea KURODA & HABE in KURODA, HABE & OYAMA, 1971,
Genus Octoporia SCARLATO & STAROBOGATOV, 1983,Genus Bathyneara SCARLATO & STAROBOGATOV, 1983,
Genus Nordonearea OKUTANI, 1985, Genus Soyomya OKUTANI, 1985, Genus Pseudogrippina MARSHALL, 2002
and Genus Krylovina VALENTICH-SCOTT, 2012.
Remark
The validity of the Genera
Naera GRAY in GRIFFITH & PIDGEON, 1834, Plectodon CARPENTER, 1865 and Pseudoneara STURANY, 1901
remains controversed.
Pictorial information
Good illustrations of some taxa of the following Genera:
Genus Cuspidaria NARDO, 1840, Genus Naera GRAY in GRIFFITH & PIDGEON, 1834,
Genus Cardiomya ADAMS, 1864, Genus Halonympha DALL & SMITH, 1886,
Genus Myonera DALL & SMITH, 1886, Genus Rhinoclama DALL & SMITH, 1886,
Genus Tropidomya DALL & SMITH, 1886, Genus Protocuspidaria ALLEN & MORGAN, 1931
and Genus Austroneara POWELL, 1937
were published by Abbott and Dance in their: Compendium of Seashells (1998).
Generalities
This Family regroups a large number of
small marine bivalves with a solid shell of 0,5 to 2,5 centimetres in length.
Some of their populations were observed in brackish environments.
The majority of its taxa inhabits coastal waters, but some of their representatives
were collected at more than 2.000 metres depth.
This Family has a world-wide distribution, but some of its specific taxa are endemic to restricted areas,
which is obvious when reading their name.
E.g.: For the generotype:
C. arctica (SARS, 1859), C. azorica (SMITH, 1855), C. capensis (SMITH, 1855),
C. chinensis (GRAY in GRIFFITH & PIDGEON, 1833), C. Formosa VERRILL in BUSH, 1898,
C. guineensis KNUDSEN, 1970, C. hawaiensis DALL, BARTSCH & REHDER, 1938
and C. kerguelensis (SMITH, 1885).
Geological records
Some fossils of Ladinian Age (Middle Triassic) are considered as their first representative taxa.
Some references for Family Cuspidariidae
ABSALAO, R. S. & DE CASTRO OLIVEIRA, C. D. 2011: The Genus Cuspidaria (Pelecypoda: Septibranchia:
Cuspidariidae) from the Deep Sea of Campos Basin, Brazil, with Descriptions of Two New Species. Malacologia. 54(2):119-138.
DALL, W. H. 1886: Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf
of Mexico (1877-78) and in the Carribean Sea (1879-80), by the U.S. Coast Survey steamer Blake, Lieut.Commander C.D. Sigsbee, U.S.N. and Commander J.R. Bartlett, U.S.N. commanding. XXIX. Report on the
Mollusca. Part 1: Brachiopoda and Pelecypoda. Bulletin of the Museum of Comparative Zoölogy at Harvard
College. 12(6): 171-318, pls. 1-9.
KNUDSEN, J. 2005: Anomalodesmata (bivalvia) from the Surinam Shelf, the Caribbean Region. Basteria. 69(46): 121-144.
-38-
OKUTANI, T. 1975: Deep-sea bivalves and scaphopods collected from deeper than 2,000 m in the northwestern
Pacific by the R. V. Soyo-Maru and the R. V. Kaiyo-Maru during the years 1969-1974. Bulletin of the Tokai
Regional Fisheries Research Laboratory. 82: 57-87, pls.: 1-3.
PRESTON, H. B. 1916: Report on a collection of Mollusca from the Cochin and Ennur backwaters. Records of
the Indian Museum. 12(1): 27-39.
SCARLATO, O. A. 1972: New species of Cuspidariidae (Septibranchia, Bivalvia) from the seas of Soviet far
east. In: MEDVEDEV, G. S. Ed. 1972: New species of terrestrial and marine invertebrates.Trudyi Zologicheskogo Instituta. 52: 121-128.
STURANY, R. 1901: Expedition S.M. Schiff Pola in das Rothe Meer. Nördliche und südliche Hälfte. 1875/96
und 1897/98. XIV. Zoologische Ergebnisse. Lamellibranchiaten des Rothen Meeres. Denkschriften der Mathematisch-Naturwissenschaftlichen Classe der Kaiserlichen Akademie der Wissenschafte. Wien. 69: 255-295, pls.:
1-7.
12.13. Family Verticordiidae STOLICZKA, 1870
(Plate 30)
Systematics
According to WoRMS 2015, this Family regroups the nine extant Genera:
Genus Verticordia de SOWERBY, 1844, Genus Trigonulina d’ORBIGNY, 1842,
Genus Laevicordia SEGUENZA, 1876, Genus Haliris DALL, 1886, Genus Halicardia DALL, 1895,
Genus Spinosipella IREDALE, 1930, Genus Vertambitus IREDALE, 1930, Genus Vertisphaeara* IREDALE, 1930
and Genus Simplicicordia KURODA & HABE in KURODA, HABE & OYAMA, 1971.
And two extinct Genera:
Genus Kurinoa MARWICK, 1942 and Genus Pecchiola SAVI & MENEGHINI, 1850.
*Spelling according to WoRMS 2015: List of the accepted Genera of the Family Verticordiidae.
Generalities
This Family regroups very small deep-waters bivalves with ribbed valves.
The Family has a world-wide distribution, but many of its representatives are endemic to some countries:
E.g.:
Verticordia australensis SMITH, 1885, Verticordia tasmanica THIELE & JAECKEL, 1931,
Halicordia nipponensis OKUTANI, 1957 and Halicordia philippinensis POUTIER, 1981.
Geological records
This Family is represented by some taxa discovered in strata of Albian Age (Cretaceous Period).
Some references for Family Verticordiidae
ALLEN, J. A. & TURNER, J. F. 1974: On the Functional Morphology of the Family Verticordiidae (Bivalvia)
with Descriptions of New Species from the Abyssal Atlantic. Philosophical Transactions - Biology. 268(894):
401-532.
DALL, W. H., 1895: Scientific results of explorations by the U.S. Fish Commission steamer Albatross. XXXIV.
Report on Mollusca and Brachiopoda dredged in deep water, chiefly near the Hawaiian Islands, with illustrations
of hitherto unfigured species from northwest America. Proceedings of the United States National Museum. 17
(1032): 675-733, pls.: 23-32.
DOGAN, A., ÖNENEN, M., ÖZTÜRK, B. & BITLIS, B. 2009: Two Rare Deep-Sea Bivalve Species from the
Levantine Coast of Turkey: Bathyarca philippiana (Nyst, 1848) and Verticordia granulata Seguenza G., 1860.
Turkish Journal of Zoology. 33: 225-230.
HEILPRIN, A. 1881: Remarks on the Molluscan Genera Hippagus, Verticordia and Pecchiolia. Proceedings of
the Academy of Natural Sciences of Philadelphia. 33: 423-428.
IREDALE, T. 1930: More notes on the marine Mollusca of New South Wales. Records of the Australian
Museum. 17(9): 384-407, pls.: 62-65.
-39-
KURODA, T., HABE, T. & OYAMA, K. 1971: The Sea Shells of Sagami Bay. Maruzen Co., Tokyo. XIX +
741 p. (Japanese text), 1-489 (English text), 1-51 (Index), pls.: 1-121.
POUTIERS, J. M. & BERNARDe, F. R. 1995: Carnivorous bivalve molluscs (Anomalodesmata) from the
tropical western Pacific Ocean, with a proposed classification and a catalogue of Recent species. In BOUCHET,
P. Ed. 1995: Résultats des Campagnes MUSORSTOM, vol. 14. Mémoires Muséum national Histoire naturelle.
167: 107-187.
12.14. Family Teredinidae RAFINESQUE, 1815 and
Family Xylophagaidae HAGA & KASE, 2013
Systematics
The systematics of these two Families were detailed in precedent paragraphs.
Taxonomic remark
WoRMS 2015 still mentions Xylophagidae PURCHON, 1941.
(No more comments)
All the extant and extinct representatives of these two Families are xylophagous. The juveniles and the adults are
protected by the wood they parasite, but the populations of all their taxa remain vulnerable during their larval
nectic stage.
12.15. Family Hiatellidae GRAY, 1824
(Plate 26)
Systematics
According to WoRMS 2015, this Family only regroups the four valid extant Genera:
Genus Hiatella BOSC, 1801, Genus Cyrtodaria REUSS, 1801, Genus Panopea MENARD de la GROYE, 1807
and Genus Panomya GRAY, 1857.
Taxonomic remark
According to WoRMS 2015, the eighteen other generic taxa proposed by diverse authors
are considered as synonyms of these ones.
Ecological remark
Except for the species included in the Genus Hiatella, all the representatives of this Family seem to have been,
such as the representatives of the Family Myidae, embedding mollusks.
The juveniles and the adults are protected by the mass of the sediments in which, since the Eocene Period, they
seem to live deeper and deeper embedded.
Genus Hiatella BOSC, 1801
Systematics
According to GOFAS, 2014, the Genus Hiatella regroups the six extant species:
Hiatella rugosa LINNAEUS, 1767, Hiatella pholadis LINNAEUS, 1767, Hiatella australis de LAMARCK, 1818,
Hiatella solida SOWERBY, 1834, Hiatella azaria DALL, 1881 and Hiatella arenacea SMITH, 1910.
According to WoRMS 2015, the Genus Hiatella regroups the five extant species:
Hiatella rugosa (LINNAEUS, 1767), Hiatella arctica (LINNAEUS, 1767), Hiatella australis (de LAMARCK, 1818),
Hiatella azaria (DALL, 1881) and Hiatella arenacea SMITH, 1910.
Remark
Differences between the List of WoRMS and other scientific Lists, are common facts
which must not disturb any searchers.
Ecology
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All the extant representatives of this Genus are nearly immobile individuals feeding on organic particles in
suspension in marine waters.
They inhabit rock cavities or cavities bored by other mollusks, such as these bored by representative taxa of the
Genus Martesia. This squatter-comportment exists, at least, since the lowermost Miocene.
This Genus is represented by species having a world-wide distribution. H. arctica is considered to be represented
by populations inhabiting Artic or Antarctic waters as well as subtropical and tropical zones, from coastal waters
to zones of more than 150 metres depth.
In subtropical and tropical zones, the extant taxa attributed to this Genus generally inhabit deeper waters than
these living in polar zones. It seems they are, more or less, isothermal invertebrates.
After a glacial period, many of their populations seem to split into two groups: One progressively inhabiting
northern more areas, the other southern more areas.
Geological records
Representative populations of the Genus Hiatella are common fossils in many north-West European strata
of Miocene and Pliocene Ages.
Genus Cyrtodaria REUSS, 1801
Systematics
According to Skaphandrus 2015, this Genus regroups the two extant species:
Cyrtodaria siliqua (SPENGLER, 1793) and Cyrtodaria kurriana DUNKER, 1861.
The diverse populations of the two sole extant representatives of this Genus
inhabit sandy bottoms in shallow marine and estuarian zones.
The populations of Cyrtodaria siliqua are scattered off the coasts of Labrador to these of Rhode Island.
(Northern West Atlantic Ocean).
The first populations of Cyrtodaria kurriana were discovered in the Sea of Beaufort
(Artic coasts of North America).
Some European geological records
Shells, with valves in connextion, of specific taxa attributed to the Genus Cyrtodaria
are common fossils of Belgian Miocene and Lower Pliocene strata.
Fossil representatives of this Genus were also collected in the Miocene of the Netherlands.
Remark
These fossils demonstrate that this Genus seems to become endemic to northern America after
the Pliocene Period.
Genus Panopea MENARD de la GROYE, 1807
(Plates 25 and 26)
Systematics
According to WoRMS 2015, the Genus Panopea regroups the ten extant species:
Panopea aldrovandi MENARD de la GROYE, 1807, Panopea glycimeris (BORN, 1778),
Panopea australis (SOWERBY I, 1883), Panopea zelandica (QUOY & GAIMARD, 1835),
Panopea abbreviata VALENCIENNES, 1839, Panopea generosa GOULD, 1850, Panopea japonica ADAMS, 1850,
Panopea bitruncata (CONRAD, 1872), Panopea globosa DALL, 1898 and Panopea smithae POWELL, 1950.
And two fossil taxa:
Panopea abrupta (CONRAD, 1849) and Panopea worthingtoni HUTTON, 1873.
Remark
-41-
This Genus is represented by numerous other fossil taxa and is fairly common in the Belgian Miocene.
Individuals of its specific taxa have been collected in muddy to sandy sea bottoms,
From coastal zones to depths of more than four hundred metres.
This Genus is widely distributed in the southwestern and the northwestern Atlantic Ocean
and in the northeastern Pacific Ocean.
Geological record
Cretaceous Period to Present Times.
Genus Panomya GRAY, 1857
Systematics
Accoording to WoRMS 2015, this Genus regroups the five extant species:
P. norvegica (SPENGLER, 1793), P. priapus (TILESIUS, 1822), P. ampla DALL, 1898,
P. nipponica NOMURA & HATAI, 1935 and P. aquaticavis TIBA, 1988.
All the populations of these taxa feed upon organic particles in suspension in the marine waters they inhabit.
These specific taxa inhabit diverse zones in the North Atlantic (E.g.: Norvegian coasts)
and in the North Pacific (E.g.: Japanese coasts).
Remark
Geologically, such a present geographical distribution is difficult to explain.
Geological record
Fossil shells attributable to this Genus were discovered in Miocene deposits of Denmark.
Some references for Family Hiatellidae
BIELER, R., CARTER, J. G. & COAN, E. V. 2010: Classification of Bivalve families. Pp.: 113-133. In: BOUCHET, P. & ROCROI, J.-P. 2010: Nomenclator of Bivalve Families. Malacologia. 52(2): 1-184.
BROWN, R. A. & THUESEN, E. V. 2011: Biodiversity of mobile benthic fauna in geoduck (Panopea
generosa) aquaculture beds in southern Puget Sound, Washington. Journal of Shellfish Research. 30: 771-776.
GORDILLO, S. 2001: Puzzling distribution of the fossil and living genus Hiatella (Bivalvia). Palaeogeography,
Palaeoclimatology, Palaeoecology. 165: 231-249.
HUNTER, W. R. 1949: The structure and behaviour of Hiatella gallicana (Lamarck) and H. arctica (L.), with
special reference to the boring habit. Proceedings of the Royal Society of Edinburgh. Biological S. 63: 271-289.
JANSEN, A. W. 1964: De E3 Scheldetunnel. Medelingen van de Werkgroep voor Tertiaire en Kwartaire
Geologie. 4: 50-54.
KANTOR, Y. I. & SYSOEV, A. V. 2005: Catalogue of Molluscs of Russia and Adjacent Countries. 627 p.
KELLY, S. R. A. 1980: Hiatella-a Jurassic bivalve squatter? Palaeontology. 23: 769-781.
RADWANSKI, A., FRIJS, H. & LARSEN, G. 1975: The Miocene Hagenør-Boerup sequence at Lillebælt
(Denmark): its biogenic structures and depositional environment. Bulletin of the Geological Society of Denmark.
24: 229-260.
SCHNEIDER, S. & KAIM, A. 2012: Early ontogeny of Middle Jurassic hiatellids from a wood-fall association:
implications for phylogeny and palaeoecology of Hiatellidae. Journal of Molluscan Studies. 78(1): 119-127.
-42-
SIGNORELLI, J. H. & ALFAYA, J. E. F. 2014: Panopea abbreviata (Bivalvia: Hiatellidae) in the southwestern
Atlantic Ocean, taxonomic revision and anatomy. Malacologia. 57(2): 279-293.
SIMONARSON, L. A. 1974: Recent Cyrtodaria and its fossil occurrence in Greenland. Bulletin of the Gelogical
Society of Denmark. 23: 65-75. PDF on-line.
STRAUCH, F. 1968. Determination of Cenozoic sea-temperatures using Hiatella arctica (Linné). Palaeogeography, Palaeoclimatology, Palaeoecology. 5: 213-233.
VADOPALAS, B., PIETSCH, T. W. & FRIEDMAN, C. S. 2010: The proper name for the geoduck:
resurrection of Panopea generosa Gould, 1850, from the synonymy of Panopea abrupta (Conrad, 1819)
(Bivalvia: Myoida: Hiatellidae). Malacologia. 52(1): 169-173.
YAZIKOVA, O. V. 2001: Upper Jurassic and Cretaceous Hiatellidae (Bivalvia) of Siberia (morphology,
variability, facies, confinement and stratigraphic range). Novosti Paleontologii i Stratigrafii, suppl. to Geologiya
i Geofizika, 4:71-81.
YONGE, C. M. 1971: On functional morphology and adaptive radiation in the bivalve superfamily Saxicavacea
(Hiatella =Saxicava), Saxicavella, Panomya, Panope, Cyrtodaria). Malacologia. 11(1): 1-44.
12.16. Family Pholadidae de LAMARCK, 1809
(Plate 22)
Systematics
This Family seems to regroup the fourteen following extant Genera:
Genus Pholas LINNAEUS, 1758, Genus Pholadidea TURTON, 1819, Genus Martesia de SOWERBY, 1824,
Genus Barnea RISSO, 1826, Genus Jouannetia DESMOULINS, 1828, Genus Zirfaea GRAY, 1842,
Genus Penitella VALENCIENNES, 1846, Genus Parapholas CONRAD, 1848, Genus Chaceia TURNER, 1855,
Genus Cyrtopleura TRYON, 1862, Genus Diplothyra TRYON, 1862, Genus Netastoma CARPENTER, 1864,
Genus Nettastomella CARPENTER, 1865 and Genus LignopholasTURNER, 1955.
Taxonomic remarks
The Genus Xylophaga TURTON, 1822, formerly included in this Family,
is now considered as the generotype of the Family Xylophagaidae HAGA & KASE, 2013.
The Genus Xyloredo TURNER, 1972 is, presently,
considered as a member of the Family Xylophagaidae HAGA & KASE, 2013.
Ecological adaptations
This Family has for singularity the regrouping of partly embedding Genera, xylophagous Genera
and lithophagous Genera.
The extant taxa of the Genus Penitella are representative species of the Pholadidae living partially embedded.
The extant taxa of the Genera Pholas and Lignopholas are representative species of the xylophagous Pholadidae.
The extant taxa of the Genus Martesia are representative species of the lithophagous Pholadidae.
Some references for Family Pholadidae
ANSELL, A. D. & NAIR, N. B. 1969: The mechanisms of boring in Martesia striata Linné (Bivalvia:
Pholadidae) and Xylophaga dorsalis Turton (Bivalvia: Xylophaginidae). Proceedings of the Royal Society of
London, Series B. 174: 123-133.
DISTEL, D. L., AMIN, M., BURGOYNE, A., LINTON, E., MAMANGKEY, G., MORRILL, W., NOVE, J.,
WOOD, N. & YANG, J. 2011: Molecular phylogeny of Pholadoidea Lamarck, 1809 supports a single origin for
xylotrophy (wood feeding) and xylotrophic bacterial endosymbiosis in Bivalvia. Molecular Phylogenetics and
Evolution. 61: 245-254.
HOAGLAND, K. E. & TURNER, R. D. 1981: Evolution and adaptive radiation of wood-boring bivalves
(Pholadacea). Malacologia. 21:111-148.
NAIR, N. B. & ANSELL, A. D. 1968: The mechanism of boring in Zirphaea crispata (L.) (Bivalvia: Pholadidae). Proceedings of the Royal Society, Series B. 170: 155-173.
SELIG, H. 1927: The kinetics of dark adaptation. The Journal of General Physiology. 10(5): 781-809.
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TAPILA, L., ROBERTS, E. M., BOUARE, M., SISSOKO, F. & O’LEARY, M. A. 2004: Bivalve Borings in
Phosphatic Coprolites and Bone,Cretaceous–Paleogene, Northeastern Mali. Palaios. 5(19): 565-573.
TURNER, R. D. 1955: The family Pholadidae in the western Atlantic and the eastern Pacific, Part II:
Martesiinae, Jouannetiinae and Xylophagainae. Johnsonia. 3: 65-160.
TURNER, R. D. & SANTHAKUMARAN, L. N. 1989: The Genera Martesia and Lignopholas in the IndoPacific (Mollusca: Bivalvia: Pholadidae). Ophelia. 30(3): 155-186.
12.17. Family Gastrochaenidae GRAY, 1840
(Plates 18 and 29)
Systematics
According to WoRMS 2015, this Family regroups the eighth valid extant Genera:
Genus Gastrochaena SPENGLER, 1783, Genus Rocellaria de BLAINVILLE, 1828,
Genus Cucurbitula GOULD, 1861, Genus Spengleria TRYON, 1861, Genus Dufoichaena EAMES, 1951,
Genus Eufistulana EAMES, 1951,Genus Lamychaena FRENEIX, 1979 and Genus Spenglerichaena CARTER, 2011.
Morphological remark
This Family regroups bivalve taxa having some of the strangest adaptations.
The taxa attributed to this Family are borers of diverse calcareous materials, such as the extern sides of septaria
or the very thick valves of some ostreid bivalves *.
*See in Géominpal Belgica. 5.2.: Individuals and colonies of Martesia peroni COSMANN & LAMBERT, 1884 discovered into
valves of Pycnodonte callifera (de LAMARCK, 1819).
Paleocological singularities
1. Succesive inhabitants
During the beginning of the sedimentation of the Sands of Edegem (lowermost marine Belgian Miocene
Formation), many septaria were extracted by submarine erosion from the upper part of the Boom Clay Member.
These ones constituted ideal anchorage places for diverse invertebrates*, but also relatively easy surfaces to bore
for individuals of Martesia peroni COSMANN & LAMBERT, 1884.
*Such as individuals of the little Anthozoa: cf. Genus Cariophylla de LAMARCK, 1801.
Michel Girardot had mentionned * to the senior-author his ancient discovery: Three to four bivalves have
successively inhabited these holes filled by fine glauconite and sand cores: one or two generations** of
Martesia peroni followed by one individual of Hiatella arctica (LINNAEUS, 1767)*** and one individual of
Coralliophila cf. lithophagella de LAMARCK, 1819***.
On the verges of the E3, at Borgerhout (Antwerp Province, Belgium), fifty-two similar associations were rediscovered in 1969 by the senior-author. A dozen of bored septaria were stored in the repository of the B..G.S.
*In September 1969.
**The second one differing from the first by its obliged smaller size and its nearly smooth valves.
***Determination: Arie Jansen, 1964.
Other common inhabitants of this sandy bottom were very large specimens of an uundetermined representative
attributable to the Genus Flabellum LESSON, 1831 (Family Flabellidae BOURNE, 1905, Hexacorallia, Anthozoa).
2. Strange constructions
Other taxa, such as the species of the Genus Cucurbitula, are builders of superposed loges constituted by walls
of fine sand cores strongly cemented (See Plate 19).
3. Shell of a Cucurbitula cucurbitula fixed on gastropod shells
See: Conchology Encyclopedia: photograph 631708 illustring
a Cucurbitula cucurbitula fixed on the shell of a small gastropod.
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The diverse extant specific taxa of this Family are common inhabitants of warm marine waters.
Geological record
In the Paris Basin, its generotype, the Genus Gastrochaena, is represented since the uppermost Eocene
by Gastrochaena ampullaria (de LAMARCK,1806).
Enigmatic concretion
(Plate 43)
This huge* concretion was stored on the right side of the main entry of the former ** Musée royal d’Histoire
Naturelle de Belgique. It is morphologically identic to the centimetric sized habitation of an extant individual of
the species Cucurbitula cymbium (SPENGLER, 1783).
*Of metric size.
**Since 1948: Institut royal des Sciences Naturelles de Belgique.
Some references for Family Gastrochaenidae
SPENGLER, L. 1783: Beskrivelse over en nye Slægt af toskallede Muskeler, som kan kaldes Gastrochæna, i tre
foranderlige Arter, hvoraf hver boer i et forskielligt Ormehuus.. Nye Samling af det Kongelige Danske Videnskabers Selskabs Skrifter. 2: 174-183, pl.: 1.
FELDER, D. L. 2009: Gulf of Mexico Origin, Waters, and Biota: Biodiversity, Volume 1. Texas A&M
University Press. 1312 p. Search: Family Gastrochaenidae.
KEEN, A. M. 1971: Sea Shells of Tropical West America: Marine Mollusks from Baja California to Peru.
Stanford University Press. XXXp. Search: Family Gastrochaenidae.
12.18. Family Clavagellidae d’ORBIGNY, 1844
(Plate 31)
Systematics
According to WoRMS 2015, this Family regroups the five Genera:
Genus Clavagella de BLAINVILLE, 1817, Genus Bryopa GRAY, 1847, Genus Dacosta GRAY, 1852,
Genus Stirpulina STOLICZCA, 1870 and Genus Dianadema MORTON, 2003.
The diverse extant and extinct specific taxa of this Family were and remain
common inhabitants of warm marine waters.
Their extant populations inhabit sandy bottoms off the coasts of the Indian and Pacific Oceans.
Geological records
The Genus Clavagella seems to be the first taxon to have fossil representatives. Its oldest specimens were
discovered in levels of Cenomanian Age in California (U.S.A.). In Europe, they appeared during the Paleocene
Age in the Franco-Belgian Basin.
In Belgium, the senior-author has collected tenths of individuals of Stirpulina cf. bacillum (BROCCHI, 1814) in a
sandstone block of the Sands of Lede Formation* (former Belgian Ledian Stage or lower part of the Upper
Eocene).
*Dejonghe Sandpit, at Meldert (Flemish Brabant, Belgium).
Some references for Family Clavagellidae
BIELER, R., CARTER, J. G. & COAN, E. V. 2010: Classification of Bivalve families. Pp.: 113-133. In: BOUCHET, P. & ROCROI, J.-P. 2010: Nomenclator of Bivalve Families. Malacologia. 52(2): 1-184.
MORTON, B. 2007: The evolution of the watering pot shells (Bivalvia, Anomalodesmata: Clavagellidae and
Penicillidae). Records of the Western Australian Museum. 24: 19-64.
MORTON, B. & GREBNEFF, A. 2011: A new species and a new record of endobenthic Clavagellidae (Bival-
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via: Anomalodesmata: Clavagelloidea) from the Oligocene and Miocene of New Zealand. New Zealand Journal of Geology and Geophysics. 54(1): 125-134.
SMITH, B. J. 1971: A review of the Family Clavagellidae (Pelecypoda, Mollusca) from Australia, with
descriptions of two new species. Journal of the Malacological Society of Australia. 2(2): 135-161.
12.19. Family Penicillidae BRUGUIERE, 1789
(Plate 31)
Systematics
According to WoRMS 2015, this Family regroups the six Genera:
Genus Brechites GUETTARD, 1770, Genus Verpa RÖDING, 1798, Genus Foegia GRAY, 1842,
Genus Humphreyia GRAY, 1858, Genus Nipponoclava SMITH, 1976 and
Genus Kendrickiana MORTON, 2004.
Principal taxonomic remarks
Fide WoRMS 2015:
The Genus Penicillus BRUGUIERE, 1789 is considered as a synonym of Genus Verpa RÖDING, 1798,
but the name Family Penecillidae BRUGUIERE, 1789 is conserved.
No comments.
The Genus Aspergilllum de LAMARCK, 1818 is considered as a synonym of Genus Brechites GUETTARD, 1770.
The Genus Clepsydra SCHUMACHER, 1817 is considered as a synonym of Genus Brechites GUETTARD, 1770.
More important
Some malacologists consider that the taxa attributed to the Families Clavagellidae and the Penicillidae
may be regrouped into a single Family, the Family Clavagellidae.
In this conception, it is a little easier to understand their Natural History.
Generalities
These bivalves have the most aberrant morphologies of the Class Bivalvia.
Their juvenile shell presents a normal morphology, but quickly their valves unit together and
initiate the building of an elongated tube presenting an upper multi-perforated, watering-pot closure.
All the extant generic taxa of this Family are endemic to some countries:
The representatives of the Genus Brechites GUETTARD, 1770 inhabit the Red Sea.
These of Genus Verpa RÖDING, 1798 inhabit some parts of the Philippines.
The diverse populations of the Genus Foegia GRAY, 1842 are scattered
along some western Australia and New-Zealand coasts.
These representatives of the Genus Humphreyia GRAY, 1858 inhabit some coasts of New-Zealand.
These representatives of the Genus Nipponoclava SMITH, 1976 inhabit some coasts of Japan,
and these of the Genus Kendrickiana MORTON, 2004 are endemic to some southern Australian coasts.
Geological records
Oligocene to Present Times.
Some references for Family Penicillidae
HUBER, M. 2010: Compendium of bivalves. A full-color guide to 3,300 of the world’s marine bivalves. A
status on Bivalvia after 250 years of research. Hackenheim: ConchBooks. 901 p.
MORTON, B. 2004:The biology and functional morphology of Foegia novaezelandiae (Bivalvia: Anomalodesmata: Clavagelloidae) from Western Australia. Malacologia. 46: 37-55.
MORTON, B. 2004: The biology and the functional morphology of Kendrickiana gen. nov. veitchi (Bivalvia:
Anomalodesmata: Clavagelloidea) from southern Australia. Invertebrate Biology. 123: 244-259.
MORTON, B. 2007: The evolution of the watering pot shells (Bivalvia, Anomalodesmata: Clavagellidae and
Penicillidae). Records of the Western Australian Museum. 24: 19-64.
SAVAZZI, E. 2005: The function and evolution of lateral asymmetry in boring endolithic bivalves. Paleon-
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tological Research. 9(2): 169-187.
TAN, S. K., TAN, S. H. & LOW, M. E. Y. 2011: A reassessment of Verpa penis (Linnaeus, 1758) (Mollusca:
Bivalvia: Clavagelloidea), a species presumed nationally extinct. Nature in Singapore. 4: 5-8.
12.20. Family Pholadomyidae KING, 1844
(Plate 28)
Systematics
According to WoRMS 2015, this Family regroups the two Genera:
Genus Pholadomya SOWERBY, 1823 and Genus Flabellomya ROLLIER, 1911.
Principal taxonomic remarks
WoRMS 2015, immediately after this assertion, mentions that
Genus Flabellomya ROLLIER, 1911 is a synonym for Genus Pholadadomya SOWERBY, 1823.
Try to understand.
The extant Genus Panacca DALL, 1905 is generally included in this Family.
Generalities
Such as their extant representatives, all the fossil taxa of this Family
were discovered in sediments deposited in shallow marine water environments.
The rare individuals of its extant representative taxon Pholadomya candida SOWERBY, 1823
were discovered in the Caribbean Sea.
Geological records
The Genus Pholadomya is represented by numerous fossil taxa discovered in strata of Lower Cretaceous Age,
on both sides of the south Atlantic Ocean (Africa and Brazil).
During the Upper Cretaceous Period,
this Genus was still represented on both sides of the north Atlantic Ocean.
The species Pholadomya candida SOWERBY, 1823, living in shallow waters of the Caribbean Sea,
seems to be its sole extant representative.
Some references for Family Pholadomyidae
CAJUS, G. D. 2012: Palaeoecology, facies and stratigraphy of shallow marine macrofauna from the Upper
Oligocene (Palaeogene) of the southern Pre-North Sea Basin of Astrup (NW Germany). Central European Journal of Geosciences. 4(1): 163-187.
CAMPBELL, H. J. & GRANT-MACKIE, J. A. 1995: Jurassic Pholadomyidae (Bivalvia) from New Zealand
and New Caledonia. New Zealand Journal of Geology and Geophysics. 38(1): 47-59.
COAN, E. V. 2000: A new species of Panacca from Chile (Bivalvia: Pholadomyoidea: Parilimyidae).
Malacologia. 42: 165-170.
DIAZ, J. M. & BORRERO, F. 1995: On the occurrence of Pholadomya Candida Sowerby, 1823 (Bivalvia:
Anomalodesmata) on the Caribbean Coast of Colombia. Journal of Molluscan Studies. 61(3): 406-408.
DIAZ, J. M., GAST, F. & TORRES, D. C. 2009: Rediscovery of a Caribbean living fossil: Pholadomya candida
G. B. Sowerby I, 1823 (Bivalvia: Anomalodesmata: Pholadomyoidea). The Nautilus. 123(1): 19-20.
JAITLY, A. K. 2013: Comments on the Middle Jurassic Pholadomyoids of Kahchch, Western India. Journal of
the Palaeontological Society of India. 58(1): 51-60.
LAZO, D. G. 2007: The bivalve Pholadomya gigantea in the Early Cretaceous of Argentina: Taxonomy,
taphonomy, and paleogeographic implications. Acta Palaeontologica Polonica. 52 (2): 375-390.
MORTON, B. 1980: The anatomy of the ‘living fossil’ Pholadomya candida Sowerby, 1823 (Mollusca:
Bivalvia: Anomalodesmata). Videnskabelige Meddelelser Fra Dansk Naturhistorik Forening I Kjøbenhavn. 142:
7-102.
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TAKENORI, S. & TAKASHI, O. 2007: A new Species of Panacca (Bivalvia: Pholadomyoidea: Parilimyidae)
from Kanesunose Bank, off Central Japan. Venus. 67(4): 372-375.
ZINMEISTER, W. J. 1978: Review of the bivalve genus Pholadomya from the Tertiary of California and the
description of two new species.Veliger. 21(2): 232-235.
13. Conclusions
13.1. Generalities
The shell of the Bivalvia includes a very large diversity of morphological and ecological adaptations, allowing
some of their taxa the common practice of lithophagy, xylophagy or blockhaus-building.
Some of these adaptations look like ecological attempts to resist against extern aggressions which could be able
to modify their genetic code, or which have modified the genetic code of their ancestors at diverse geological
periods.
13.2. Concerning the systematic position of the extant and extinct teredinid taxa
All the extant and, as far as known, all the extinct representatives of all the specific taxa attributed to the two
Families Teredinidae and Xylophagaidae have the following characteristics in common.
They possess a shell with two identic calcareous valves of reduced sizes, an elongated styloid apophyse *, an
extremely elongated siphon and one pair of biomineralised organites called paletts. They feed upon drifting
pieces of wood, mangrove roots or decayed vegetal remains which means they are all strictly lithophagous.
*Intern narrow calcareous extension into the shell serving for the fixation of the pedal muscle.
As far as known, their DNA presents strong affinities.The number of eggs produced by their females may rise up
to one million. Their larvae are nectic and ensure a large distribution for the majority of their extant taxa.
Many of their taxa have an oceanic, if not world-wide geographical distribution and the geographical dispersion
of some of their taxa endemic to restricted areas may result from human maritime trades*.
*References to the Portuguese and Spanish explorations, but also to the Phoenicians colonisations of the Atlantic coasts of
southern Spain and northern Morocco in the Antiquity.
All these common singularities seem to be sufficient for the senior-author, to propose a regrouping of these two
Families in a distinct Order: Order Terediniformes Order nov.
The possession of an elongated styloid apophyse allows supposing that their ancestors were Pholas-like bivalves
and that the Genus Martesia is one of their nearest phylogenetic parents.
In this conception, the Families* regrouping taxa being not in possession of a styloid apophyse may be considered as their farest phylogenetic parents.
*Such as the Families Corbulidae, Hyatellidae, Lyonsiidae, Myidae, Pandoridae, Pholadomyidae, Thraciidae and Veticorbulidae.
13.3. Concerning the systematic position of the fossils
discovered in the Sint-Niklaas Phosphorite Bed
Even if the morphology of the tubes of the Terediniformes is not a sufficient specific or generic criterion, it
seems obvious that all the teredinid tubes discovered in the petrified pieces of wood concentrated in this level
were not built by individuals of a single species.
Some of these tubes have a regular cylindrical shape1, others present bulbous parts2 and other ones3 have had
their growth marked by obvious strentheninghs.
1. Specimen illustrated on Plate 1
Such regular cylindrical tubes are the result of the building activity of plenty of extant representatives of the large majority of the Terediniformes.
Proposal
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Consequently, for this type of tube, the senior-author proposes:
Order Terediniformes, Family Teredinidae, sub-Family indet., Genus indet. and species indet.
Such cylindrical tubes, constituted by a succession of lightly inflated parts, may result of the building activity of
some extant representatives of the Genus Nototeredo (See: Plate 11).
Proposal
For this type of tube, the senior-author proposes:
Order Terediniformes, Family Teredinidae, sub-Family Bankiinae, cf. Genus Nototeredo, species indet.
The single extant Genus of the Family Teredinidae, as well as its American Paleocene representatives *, building tubes with a circular cross-section and some local over-thickness are attributed to the Genus Nototeredo**.
*See TURNER, S. 1966: p.15: fig.3: A, F and G.
**Taxon attributed to the sub-Family Bankiinae (Family Teredinidae).
In western Europe, the presence of the sub-Family Bankiinae is attested by some fossil reprensentatives of the
Genus Bankia, discovered in strata of Lutetian Age (Middle Lutetian) at Vandancourt (France).
Proposal
Consequently, for this type of tube, the senior-author proposes:
Order Terediniformes, Family Teredinidae, sub-Family Bankiinae, Genus Nototeredo, species indet.
4. Future investigations
These fossil remains were transmitted to Dr. Olev Vinn (University of Tartu, Estonia) for further examinations.
13.4. Concerning the present conception of the generic taxa
of the extant Myoida and Anomalodesmata
A short look on the plates illustring the variability of the morphology of the shell of some taxa attributed to some
Genera, included in the Families Corbulidae, Myochasmidae and Pandoridae, must be sufficient to understand
that the majority of their extant generic representatives require a complete systematic revision.
The research for illustrations of the diverse species presently included in one extant Genus in the Shell
Encyclopedia (www.conchology.be) furnish plenty other examples of these incoherent generic conceptions.
13.5. Concerning the present conception of the Order Anomalodesmata
The Order Myoida and the new Order Terediniformes, such as proposed, seem to constitute two homogenous
bivalve groups, but the Order Anomalodesmata looks like a melting-pot without any phylogenetic signification.
DNA analyses could certainly confirm this supposition, but a careful examination of the evolution of their larval
representatives would be sufficient and less expensive.
Some references for Anomalodesmata
mollusks from Baja California to northern Peru. 2 vols, 1258 p.
MORTON, B. 2002: Biology and functional morphology of the watering pot shell Brechites vaginiferus
(Bivalvia: Anomalodesmata: Clavagelloidea). Journal of Zoology. 257: 545-562.
MORTON, B. 2007: The evolution of the watering pot shells (Bivalvia, Anomalodesmata: Clavagellidae and Penicillidae). Records of the Western Australian Museum. 24: 19-64.
SPRY, J. F. 1964: The sea shells of Dar es Salaam: Part 2: Pelecypoda (Bivalves). Tanganyika Notes and
Records. 63.
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PLATES
Plate 1 to Plate 8
Fossil teredinid specimens
discovered in the
Sint-Niklaas Phosphorite Bed
Plate 9 to Plate 41
Xylophagous living taxa.
Living teredinids and their environments.
Order Myoida and Order Anomalodesmata.
Other Belgian Cenozoic pieces of wood
bored by fossil teredinids.
Plate 42
The largest enigmatica
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Plate 1: S.V.K. Clay Pit 4: B.G.S. Archives: N°: 42 W 394:
Oligocene: Sint Niklaas Phosphorite Bed: Piece of wood hosted by Teredo-like bivalves:
1: Detail of the tranversal section of a piece of wood presenting its central part, seasonal growth rings,
the cross section of thirteen tubes of teredinid Bivalvia and the sediments filling these ones.
Estimated diameter of this branch: 6,5 centimetres.
Photographer: Hugues Doutrelepont. See comments.
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1: View of one side of a radial section of a silicified wood-piece presenting
numerous tubes of an undetermined teredinid Bivalvia.
Height of the wood-piece: 11,5 centimetres. Photographer: Hugues Doutrelepont.
See comments.
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1a to 1c: Magnifications of three parts of the wood-piece illustrated on Plate 2.
See comments.
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1: Part of a silicified piece of wood presenting numerous tubes
of an undetermined teredinid Bivalvia.
Height of this piece of wood: 8,5 centimetres. Photographer: Hugues Doutrelepont.
See comments.
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1: Magnification of the central part of the photograph of the piece of wood illustrated on Plate 4.
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1a: Magnification of the lower extremity of another piece of wood.
1b: Magnification of the upper extremity of another piece of wood.
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1a: Complete view of this piece of wood. Height of the wood-piece: 14,6 centimetres.
1b: Detail of the lower part of the same piece of wood.
See comments.
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1a: Detail of the upper part of the same piece of wood illustrated on Plate 7.
1b: Detail of the cross section of the piece of wood illustrated on Plate 7.
On this magnification, the wood fibers are perfectly visible and identifiable.
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Plate 9: Crustacea (Isopoda, Family Limnoriidae) and Mollusca (Bivalvia, Family Teredinidae):
1: Adult male and female of Limnoria quadripunctata (MENZIES, 1957) - Gulf of Morbihan, France.
2: Nearly adult individual of Limnoria quadripunctata (MENZIES, 1957), boring in a wood piece of a pile.
3: Shell of and cast of the tube of one juvenile individual of Teredo navalis LINNAEUS, 1758.
4 and 5: Pieces of wood perforated by colonies of Teredo navalis LINNAEUS, 1758.
Common source: www.wikipedia.org . See comments.
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Plate 10: General morphology of the shell and the tube of the extant Terediniformes:
1: Tubes of a Teredinidae discovered in an immerged piece of wood.
Their extremely elongated bodies are protected by a calcareous tube they have secreted around their body.
2: View of the anterior extremity showing the small bivalve shell, the builder of this tube.
3: View of the shell. Common source: Auguste Le Roux. See comments.
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Plate 11: Family Teredinidae: Genera Bankia, Kuphus and Nototeredo:
1: Views of the valves and one pallet of an individual of Bankia carinata (GRAY, 1827)
2: Side view of the tube of a senile individual of Kuphus sp. Length: 164 centimetres.
3: View of the extremity of the tube of a juvenile individual of Nototeredo norvegica (SPENGLER, 1792)
4: Views of two irregular tubes of Nototeredo norvegica (SPENGLER, 1792)
Diverse sources. See comments.
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1
2
3
Plate 12: Morphological variability of thhe tubes of the Genus Kuphus
1. Kuphus sp. Origin: Mactan Island (The Philippines). Depth: Between 10 and 35 metres.
Uncomplete tube. Fragment’s size: 62,3 millimetres.
2. Kuphus sp. Origin: Mactan Island (The Philippines). Depth: 8 metres.
Nearly complete tube. Size: 187 millimetres.
3. Kuphus polythalamia (LINNAEUS, 1758) Origin: Coron (The Philippines). Depth: 8 metres.
Nearly complete tube. Size: 240 millimetres.
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Plate 13: Some feeding sources of the Terediniformes:
1: Rhizophora mangle LINNAEUS, 1753: A common mangrove tree with its adventive roots.
Origin: Southern Florida (U.S.A.).
2: Massive accumulation of stranded parts of dead wood, already hosted
or shortly before to be hosted by teredinid bivalves.
Origin: Eastern Pacific coast, California (U.S.A.).
Common source: www.wikipedia.org See comments.
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Plate 14: Mangrove trees of which parts of roots are commonly hosted by Teredo-like Bivalvia:
1: A very old specimen of Bruguiera cf. gymnorrhiza (LINNAEUS, 1752). Locality: coast of Bali.
2: View of an isolated Avicennia marina (FORSKALL, 1784). Locality: off southern Sri-Lanka.
Common source: www.wikipedia.org
See comments.
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Plate 15: Family Teredinidae RAFINESQUE, 1815 and Family Hiatellidae GRAY, 1824.
1: Family Teredinidae: Genus Teredo: Teredo navalis LINNAEUS, 1758.
2: Family Hiatellidae: Genus Hiatella: Hiatella arctica LINNAEUS, 1767.
3: Family Teredinidae: Genus Kuphus: 3a to 3f: Diverse tubes of Kuphus sp.
Courtesy of Guido and Philippe Poppe.
See comments.
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Plate 16: Hosts of empty tubes, Mycelium and siphons:
1 and 2: Teleostei commonly inhabiting tubes of dead Teredinidae: Family Microdesmidae (Perciformes):
1: Oxymetopon cyanoctenosum KLAUSEWITZ & CONDE, 1981. 2: Nemateleotris magnifica FOWLER, 1938.
3: Mycelium damaging a piece of wood before its drift. 4: View of one pallet of a juvenile, of one pallet of an
adult and of one pallet of a senile Kuphus sp.
5: Siphons of a colony of Kuphus sp., sub-Family Kuphinae, Family Teredinidae.
Common source: www.wikipedia.org See comments.
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Plate 17: Examples of Habitat:
1: Camouflage of Cucurbitula cymbium (SPENGLER, 1783), an Anomalodesmata.
Origin: The Philippines. Courtesy of Guido and Philippe Poppe.
2: Colony of Lithophaga lithophaga (LINNAEUS, 1758), a Mytiloida.
Origin: Croatia, Pag Island, off Simuni.
Photographer Jiri Novak. Source: www.biolib.cz
See comments.
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Plate 18: Family Teredinidae and Family Gastrochaenidae
1a to 2b: Family Teredinidae (Terediniformes): Genus Uperotus GUETTARD, 1777:
Uperotus nucivora (GMELIN, 1791).1a: View of the pair of pallets. 1b: View of the two valves in living
position. 1c: Intern and extern views of a left valve of the shell. 2a: Lateral view of a colony. 2b: Upper view of
the same colony. 3: Family Gastrochaenidae (Anomalodesmata):: Genus Gastrochaena GRAY, 1840:
Gastrochaena cuneiformis SPENGLER, 1783: Lateral view of one valve of an adult individual.
Courtesy of Guido and Philippe Poppe. See comments.
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Plate 19: Morphology of the shell of diverse borer bivalves:
1: Extern view of the two valves of Teredo navalis LINNAEUS, 1758.
2: Side view of the right valve of a Lithophaga truncata (LINNAEUS, 1758). Source: www.wikipedia.org
3: View of the left side of the shell of Jouannetia cumingii (SOWERBY II, 1849).
4: Side view of the right valve of a Hiatella anophtelma RUFFO, 1957.
5: Side view of the right valve of a Lithophaga teres (PHILIPPI, 1846).
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Plate 20: Bivalvia - Family Myidae de LAMARCK, 1809 - Genus Mya LINNAEUS, 1758:
1: Side view of an adult individual of Mya truncata LINNAEUS, 1758:
Size: 90,1 millimetres. Origin: 60 metres depth. Silver Pit, North Sea.
2: Side view of the left valve an adult individual of Mya arenaria LINNAEUS, 1758.
Size: 88,7 millimetres. Origin: Off the coasts of Lancashire (Great-Britain).
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Plate 21: Morphology of the shell of some Genera of the Order Myoida:
1: Family Thraciidae: Genus Thracia: Thracia myopsis MÜLLER, 1842.
Width: 16,3 milimetres. Origin: Finmark (Norway).
2: Family Cuspidariidae: Genus Cuspidaria: Cuspidaria nobilis (ADAMS, 1854):
Width: 49,2 milimetres. Origin: Off Alguay Island (Philippines). Depth: circa 100 metres on sandy bottom.
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Plate 22: Order Myoida: Family Pholadidae: Variability of the morphology of its generic taxa:
1a-1b: Intern and extern sides of the valves of a Barnea candida (SOULEYET, 1843).
Length: 72 millimetres. Origin: Phuket Island. Thailand.
2a-2b: Extern and intern sides of the valves of a Pholas dactylus LINNAEUS, 1758.
Length: 61 millimetres. Origin: Mariakerke Beach (Western Flanders, Belgium).
Photographer Eric Vanderhoeft. See comments.
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Plate 23: Fossil and living representatives of the Genus Pholadomya SOWERBY, 1823:
1: View of the fourth individual of Pholadomya candida SOWERBY, 1823 ever found.
Origin: Carribean Sea, off Santa Marta, Columbia. Source: See comments.
2: View of an adult individual of the fossil taxon Pholadomya puschii GOLDFUSS, 1840.
Origin: Astrup, in a level of Oligocene Age, Germany.
See comments.
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Plate 24: Variability of the morphology of shells attributed to the Family Monochamidae:
1: Monochama anomioides STUTCHBURY, 1830: Size: 23 millimetres. Origin: off Tasmanian coasts.
2: Monochama tasmanica (WOODS, 1876): Size: 17,5 millimetres. Origin: Melville Island (northern Australia).
3: Monochama anomiodes STRUCHBURRY, 1830: Size: 23 millimetres. Origin: Off Tasmanian coasts .
1 to 3: Courtesy of Guido and Philippe Poppe.
See comments.
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Plate 25: Morphology of the shell of taxa of the Families Hiatellidae and Myidae:
1: Family Hiatellidae: Genus Panopea MENARD de la GROYE, 1807: Panopea generosa GOULD, 1850:
Origin: off Hong-Kong (China). Source: www.wikipedia.org
Length of the shell: 11,6 centimetres. Photographer: Patrick Fischer.
2: Family Myidae: Genus Mya LINNAEUS, 1758: Mya arenaria LINNAEUS, 1758:
Length of the shell: 10,8 centimetres. Courtesy of Guido and Philippe Poppe.
See comments.
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Plate 26: Morphology of the shell of some representatives of the Order Myoida:
1: Genus Sphenia TURTON, 1822: Sphenia hatcheri PILSBRY, 1899.
2 : Genus Panopea MENARD de la GROYE, 1807: Panopea generosa GOULD, 1850.
3: Genus Hiatella BOSC, 1801: Hiatella arctica (LINNAEUS, 1767).
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Plate 27: Difference of the morphology of shells attributed to the Genus Corbula:
1: Corbula fortisulcata SMITH, 1879. Size: 20,4 millimetres.
Origin: The Philippines.
2: Corbula caribbea d’ORBIGNY, 1842. Size: 9,6 millimetres.
Origin: Argentina.
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Plate 28: Family Gastrochaeniidae:Genus Gastrochaena SPENGLER, 1783:
1: Lateral view of the rigth valve of a Gastrochaena plicatilis (DESHAYES, 1855):
Size: 2,7 centimetres. Origin: off Bohol, the Philippines.
2: Lateral view of the rigth valve of a Gastrochaena cuneiformis SPENGLER, 1783:
Size: 1,7 centimetres. Origin: extracted from coral in the Rynie Park, South-Africa.
3: : Lateral view of the rigth valve of a Gastrochaena dubia (PENNANT, 1777)
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Plate 29: Morphological variability of the shells attributed to the Family Pandoridae:
1: Freanamya ceylanica (SOWERBY I, 1835). Size: 25 mm. Origin: Algoa Bay, South-Africa.
2: Pandora gouldiana DALL, 1896. Size: 20,9 mm. Origin: Jefferson Port (U.S.A.), circa 5 metres depth.
3: Pandora dissimilis SOWERBY III, 1894. Size: 37,4 mm. Origin: Algoa Bay, South-Africa.
4: Pandora pinna (MONTAGU, 1803). Size: 7,5 mm. Italy: off Anzio, South of Rome, at 200 metres depth.
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Plate 30: Variability of the morphology of shells attributed to the Family Verticordiidae:
1: Euciroa millegemmata KURODA & HABE, 1952: Size: 17,6 millimetres. Origin: Algoa Bay (the Philippines).
2: Euciroa elegantissima (DALL, 1881): Size: 24,4 millimetres. Origin: Off Florida coast (USA).
3: Acreuciroa rostrata (THIELE & JAECKEL, 1931):
Size: 47 millimetres. Origin: 400 metres depth, off eastern China.
1 to 3: Courtesy of Guido and Philippe Poppe. See comments.
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Plate 31: Some extreme adaptations of the shell of Anomalodesmata:
1: Brechites philippinensis (CHENU, 1843). Height: 20,8 centimetres.
Origin: Western Australia.
2: Cucurbitula cucurbitula (SPENGLER, 1783). Height: 29 millimetres.
Origin: Andaman Sea - Thailand.
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Plate 32: Other Belgian fossil piece of wood hosted by Teredo-like bivalves:
Locality: Albert II Dock, Doel (Beveren), Eastern Flanders.
Stratigraphy: Basal Grind of the Sands of Kattendijk Formation (Lower Belgian Pliocene).
1: Intern view of a drifted piece of wood. Total length: 20,5 centimetres.
1a: Magnification of one of its hosts, classically determined as: Teredo navalis LINNAEUS, 1758.
2: Extern view of another drifted piece of wood. Total length: 17,5 centimetres.
Photographs Eric Vanderhoeft. See comments.
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1
Plate 33: DOEL - ALBERT II Dock - Top of the Sands of Merksem Formation:
Uppermost Belgian Pliocene.
1: Lot of Corbula gibba OLIVI, 1792.
Photographer: Eric Vanderhoeft. See comments.
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1
Plate 34: DOEL - ALBERT II Dock - Top of the Sands of Merksem Formation:
Uppermost Belgian Pliocene.
1: Incrustations and content of diverse Corbula gibba OLIVI, 1792.
Photographer Eric Vanderhoeft. See comments
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1
Plate 35: E3 Tunnel: Base of the Edegem Sands Formation:
1: Small septaria reworked from the top of the Boom Clay Member: Extern view.
Size: 14,2 centimetres of width.
Photographer: Eric Vanderhoeft. See comments
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1
Plate 36: E3 Tunnel: Base of the Edegem Sands Formation:
1: Half a part of a small septaria reworked from the top of the Boom Clay Member.
One of its mollusks borers, Martesia peroni COSSMANN & LAMBERT, 1844 is visible.
Heigth: 11,5 centimetres.
Photographer: Eric Vanderhoeft. See comments
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1
2
3
4
5
6
Plate 37: Martesia peroni COSSMANN & LAMBERT, 1844:
1 and 4: Two views of one specimen with a part of its extern protection.
2 and 5: Two individuals perfectly preserved.
3: Intern view of a right valve.
6: Left view of another individual.
1 to 6: Photographer Eric Vanderhoeft. See comments.
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1
2
3
4
Plate 38: Second inhabitant which occupied some holes
bored by the Martesia peroni COSSMANN & LAMBERT, 1844:
1 and 2: Intern and extern views of its left valve.
3 and 4: Extern and intern views of its right valve.
1 to 4: Photographer Eric Vanderhoeft. See comments
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Plate 39: Other Belgian fossil piece of wood hosted by Teredo-like bivalves:
1: Cross section of a part of the drifted trunk of a palm tree-like vegetal.
Total length: 22,4 centimetres.
Locality: Lambrechts Sandpit, Neder-Okkerzeel, Flemish Brabant, Belgium.
Stratigraphy: Fossil discovered at the top of the Brussels Sands Formation (Lower Lutetian).
Photograph Eric Vanderhoeft. See comments.
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Plate 40: Other Belgian fossil piece of wood hosted by teredinid-like bivalves:
1: Longitudinal section of a part of the drifted trunk of a palm tree-like vegetal.
1a: General view. Total length: 24,6 centimetres. 1b: Magnification of its lower part.
Stratigraphy: Fossil discovered at the top of the Brussels Sand Formation (Lower Lutetian).
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Plate 41: Other Belgian fossil piece of wood hosted by teredinid-like bivalves:
1a and 1b: Two magnifications of the specimen illustrated on Plate 40.
Stratigraphy: Fossil discovered at the top of the Brussels Sands Formation (Lower Lutetian).
.
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Plate 42: Largest Belgian enigmatic concretion of unknown Age and Origin:
1a: Side view. Height: 125 centimetres.
1b: Lower view: Diameter: 88 centimetres.
1c: Upper view. Magnification: x 2.
Repository: Public Parking I.R.S.N.B. (Brussels, Belgium). Photographer Eric Vanderhoeft.
See comments.
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15. Comments on the Plates
Plate 1 to Plate 8
These plates regroup photographs of all the fossil remains considered as teredinid remains
discovered, between 1983 and 2004, in the sifting residues of the Sint Niklaas Phosphorite Bed,
in its type locality: The S.V.K. Clay Pit N° 4, at Sint-Niklaas (Eastern Flanders, Belgium).
One of the important taphonomical problematic is the estimation of the origin of these pieces of wood,
another is the estimation of the duration they remained in marine waters.
Their attribution to the Gymnospermales or the Angiospermales is a proposition
made by Hugues Doutrelepont (Carpologist, M.R.A.C., Tervuren, Belgium)
and, amazingly, confirmed and precised by the gardner of the senior-author.
This gardner has studied Ebenistry and Marquetry,
but for political reasons was forced to move away from his country.
Comments on Plate 1
This piece of wood is supposed to be a fragment of a branch of a Gymnospermales of the Order Pinales. The
specific taxon to which attribute this branch is difficult to ascertain without fine slides, but could be a coastal tree
of the Genus Pinus.
Trees of the Genus Pinus are abundant and common along all the coasts having a cold to warm temperate climate.
Its seasonal growth rings are particularly obvious. The sediment filling the tubes is constituted by very fine sand
and glauconite cores. The beginning of the oxysation of the iron sulphides is responsible for their greyish coloration.
The tubes present a remarkable sub-circular section of nearly the same diameter what suggests their builders had
the same age.
Comments on Plate 2
This relatively large piece of wood only presents parts of five tubes what is not normal for pieces of wood
having drifted more than one week. But, considering that this piece of wood is a transversal part of a thick
branch, the concentration of individuals teredinids is similar to the concentration observed in the first one.
These tubes present smooth walls with numerous constriction marks. But these ones could also be considered as
successive modifications of the tunneling direction.
This phenomenon is explainable by the successive changing of positions of a piece of wood immerged and
balloted in marine waters, since a few days.
The formation of the siderite concretion is considered as posterior to the drifting period.
Comment on Plate 3
These magnifications offer better views of diverse parts of the tubes illustrated on Plate 2.
Comments on Plate 4
This photograph is a magnification of another part of another piece of wood showing a teredinid tube avoiding
what seems to have been a node of the wood *
*Generally, the place of the insertion of a small branch.
The thickness of the wall of this cylindrical teredinid tube and its sedimentary filling are perfectly visible.
Comment on Plate 5
This plate presents a higher magnification of the central part of the preceding plate. If what seems to have been a
node, really a node was, it was the insertion place of a branch with a diameter of a maximum of 15 mm.
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The very high number of inflations of the teredinid tubes allows supposing that the wood encircling this node
very hard was. This hardness could imply that this branch was not the first to grow at this place.
Comments on Plate 7
This plate presents another elongated piece of wood with teredinid tubes. Some individuals have bored this one
following the fibers of the wood, but others have pierced all its layers.
On the wall of one of the tubes which follows the wood fibers, a hole is visible what means that it remained
more or less empty.
All the tubes having pierced all the layers of this piece of wood are filled by sediments.
Comment on Plate 8
This plate presents magnifications of two parts of the piece of wood illustrated on Plate 7.
Comment on Plate 9
Damages caused by the crustacean xylophagous isopod taxa and these caused by the molluscan xylophagous
teredinid taxa have similar dramatic consequences for building and naval human activities.
Pillars of wharfs and habitations or wood-ships were, and remain, very quickly destroyed by the boring activities
of both groups.
But, the damages caused on harbor constructions always were less dramatic than these caused on naval
constructions.
Comments on Plate 10 to Plate 18
These plates regroup photographs concerning the general morphology of the type species of the Genus Teredo,
genotype of the FamilyTeredinidae,different morphologies of their tubes, pieces of wood bored by their extant
representatives, some sources of wood they bore, interesting parts of their anatomy, mycelium present on dead
woods and some inhabitants of tubes of dead teredinids.
Comments on Plate 10
(Figure 1)
The growing of the tubes, illustrated on this Plate, presents numerous, regularly spaced, interruptions. Such interruptions have been interpreted as a result of an annual growing cycles or, by other authors, as a result of a moonsoon growing cycles.
These two hypothesis ignore the frequency the ancient sailors were obliged to change some essential pieces of
their ship, completely injured by ship-worms.
Some biologists have suggested that these growing interruptions could be attributed to a moon cyclus.
Even if, it remains difficult to explain how a moon cyclus could be responsible of this phenomenon, force is to
admit that this one better explains the frequency of the hull-cleaning operations of the ancient sailors.
Additionally, this interpretation is in accordance with the data concerning the growing rate of the majority of the
extant representatives of the Genus Teredo.
(Figure 2)
This photograph suggests that the extant representatives of the Genus Teredo
are able to bore wood in diverse orientations
without to be influenced by the structure of the wood.
(Figure 3)
This photograph illustrates the thin and perfectly symmetric shell
with the first centimetre of its calcareous protective tube.
Some references
BLUM, H., F. 1922: On the effect of low salinity on Teredo navalis. University of California, Publications in
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Zoology. 22(4): 348-368.
PALVAST, P. & VAN der VELDE, G. 2011: Distribution, settlement, and growth of first-year individuals of the
shipworm Teredo navalis L. (Bivalvia: Teredinidae) in the Port of Rotterdam area, the Netherlands. International
Biodeterioration and Biodegradation. 65(3): 379-388.
Impact of the Terediniformes on human naval activities
Before Humanity was able to build metallic hulls, the hull of the wood-ships was constantly and dramatically
bored by teredinids molluscs; origin of their vernacular name: shipworms. This problem obliged all the captains,
legal or pirates, to found secure places where the cleaning of the hull and the replacement of its to damaged
elements possible was.
But all the secure bays were not in possession of trees from which the resistance was this required to furnish
some naval master-pieces. For all the marine trade-companies, the domination of some islands by the crown they
depended became a priority.
Historic reference
WOODARD, C. 2008: The Republic of Pirates, being the True and Surprising Story of the Caribbean Pirates
and the Man Who Brought Them Down. Marine Books. Houghton Mifflin Harcourt. New-York. 383 p.
ISBN: 978-0-15-101302-9.
Importance of the Mycelium
It is the lace-like and kilometric-sized mycelium-net which ensures the communication between all the specific
taxa of trees and allows transmitting warnings about potential animal aggression’s.
The mangrove trees are unable to transmit such warnings because of their lacking of a mycelium warning
system. Mycelium cannot survive in any type of salt-waters, so low it is.
The Teredinidae bore drifting pieces of dead wood and roots of mangrove trees.
If stranded more than a sea-tide, the majority of them dye.
Some references for Salinity and mangroves
CINTRON, G., LUGO, A., E., POOL, D., J. & MORRIS, G. 1978: Mangroves of Arid Environments in Puerto
Rico and Adjacent Islands. Biotropica. 10(2): pp. 110-121.
CRUZ, M.,TORRES, G. & VILLAMAR, F. 1987: Estudio de los moluscos perforadores de la madera
Rhyzophora harrisonii (Mangle) en la costa Ecuatoriana. Acta Oceanografica del Pacifico. 4(1): 121-160.
LIANG, S., ZHOU, R, C., DONG, S. S. & SHI, S. H. 2008: Adaptation to salinity in mangroves: Implication on
the evolution of salt-tolerance. Chinese Science Bulletin. 53(11): 1708-1715.
Inhabitants of empty tubes of Teredinidae
The juveniles of many species of the Family Microdesmidae R EGAN, 1912 (Order Perciformes, Teleostei, Pisces)
are commonly encountered in tubes of dead Teredinidae and supposed to feed, partially, upon their decaying remains.
The senior-author has observed three juveniles of Entelurus aequereus (LINNE, 1758) in one tube of a dead
Neoteredo sp. in a piece of wood stranded at Newport (Western Flanders, Belgium)
He also observed small colonies of isopods and small decapods in Senegal and in Bahia California (Mexico).
Reference
HENDY, I. W., EME, J., DABRUZZI, T. F., CONNOLLY, H. F., NEMBHARD, R., CRAGG, S. M. & BENNETT, W. A. 2013: Dartfish use teredinid tunnels in fallen mangrove wood as a low-tide refuge. Marine Ecology Progress Series. 486: 237-245.
:
The figure 1 allows understanding the tridimensional deformations necessary to jump from
the morphology of the shell of a Myoida to the morphology of the shell of a Terediniformes.
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The figure 2 illustrates one of the hugest tube of Nototeredo norvegica known.
The figure 3 shows the paletts at the distal extremity of the tube of an individual of Nototeredo.
The figure 4 presents a photograph of one individual of Nototeredo norvegica presenting a bifid tube.
This anomaly could be interpreted as a genetic reminiscence of the original paired siphons, characteristic for all
the extant and extinct representatives of the Order Myoida.
This plate illustrates the very large variability of the morphology of the tubes of extant specific taxa of one
teredinid Genus.
Its aim is to convince paleontologist colleagues of the non-sense to continue to give a specific name to fossil
specimens lacking of their pallets.
(Figure 1)
Example for local accumulation of drifted pieces of dead wood. If going back in marine waters, these pieces of
partly degraded wood are ideal feeding sources for any taxon of the Family Teredinididae.
(Figure 2)
Mangrove wood is another important source of food, but only for some specific taxa of the Family
Teredinididae.
But, if stranded more than a sea-tide, the majority of them dye.
Some references for Salinity and mangroves
CINTRON, G., LUGO, A., E., POOL, D., J. & MORRIS, G. 1978: Mangroves of Arid Environments in Puerto
Rico and Adjacent Islands. Biotropica. 10(2): pp. 110-121.
LIANG, S., ZHOU, R, C., DONG, S. S. & SHI, S. H. 2008: Adaptation to salinity in mangroves: Implication on
the evolution of salt-tolerance. Chinese Science Bulletin. 53(11): 1708-1715.
This plate illustrates two other forms of mangrove environments.
Only some taxa of the Order Terediniformes are able to feed upon roots of these ones.
The figure 1 illustrates the two valves and one palett of a specimen attributed to the Genus Teredo.
The figure 2 presents an oblique view of an individual of Hiatella arctica (LINNAEUS, 1767).
The figures 3a to 3f illustrate the variability of the orientation of tubes of Teredo
in diverse Philippine pieces of mangrove wood.
The figures1 and 2 illustrate two common fish-skaters of empty teredinid tubes.
The figure 3 illustrates a lace-like net of Mycelium which may reduce
the hardness of wood pieces before drifting.
But when lace-like nets of Mycelium are present, the communications between living trees are impossible.
The figure 4 shows one palett of a juvenile, an adult and senile Kuphus sp.
The figure 5 illustrates the siphons of a colony of Kuphus sp.
This plate has for principal justification the illustration of a colony of lithophagous mytilid Bivalvia.
The Order Myoida STOLICZKA, 1870 regroups all the extant and extinct xylophagous bivalves, but, the Order
Mytiloida de FERUSSAC, 1822 includes some lithophagous taxa, such as these regrouped into the Genus
Lithophaga RÖDING, 1798.
The illustration of the morphology of the shell of a typical lithophagous Mytiloida seemed necessary to better
understand the convergence of the morphologies resulting from similar habitat’s adaptation.
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Some references for lithophagous Bivalvia
HARPER, E. M., TAYLOR, J. D. & CRAME, J. A. Eds. 2001: The Evolutionary Biology of the Bivalvia.
(Geological Society of London Special Publications).
KLEEMAN, K. H. 1994: Mytilid bivalve Lithophaga in Upper Triassic coral Pamiroseris from Zlambach Beds
compared with Cretaceous Lithophaga alpina. Facies. 30: 151-154.
SMITH, S. D. A. 2011: Densities of the endolithic bivalve Lithophaga lessepsiana (Vaillant, 1865) in
Pocillopora damicornis, Solitary Islands Marine Park, northern NSW, Australia. Molluscan Research. 31(1): 4246.
This plate illustrates the difference of the morphology of a shell of the Genus Uperotus GUETTARD, 1770, a typical extant Terediniformes and the morphology of a shell of the Genus Gastrochaena SPENGLER, 1783, a typical
extant Anomalodesmata.
These plates illustrate the morphology of the shell of some generic taxa
of the Orders Myoida and Anomalodesmata.
The present systematic conception of the two Orders Myoida and Anomalodesmata
remains the subject of many controversies.
In the Order Myoida itself, the conception of the Genus Mya LINNAEUS, 1758 itself is controversable. Mya
arenaria LINNAEUS, 1758, its generotype has a bivalve shell which is not truncated, but Mya truncata LINNAEUS,
1758 has a truncated bivalve shell.
E.g.: If discovered isolated, the valves of Mya truncata LINNAEUS, 1758 may easily be attributed to the Genus
Myochama STUTCHBURY, 1830, the generotype of the Family Myochamidae C ARPENTER, 1861.
The astonishing diversity of morphologies of the generic taxa attributed to the Order Anomalodesmata makes
difficult to believe in its homogeneity.
Some references concerning some taxa of the Order Anomalodesmata
HARPER, E. M., DREYEr, H. & STEINER, G. 2006: Reconstructing the Anomalodesmata (Mollusca: Bivalvia): morphology and molecules. Zoological Journal of the Linnean Society.148: 395-420.
PREZANT, R. S., SUTCHARIT, C., CHALERWAHT, K., KAKHAI, N.,DUANGDEE, T. & and
DUMRONGROJWATTANA, P.. 2008: Population study of Laternula truncata (Bivalvia: Anomalodesmata:
Laternulidae) in the mangrove sand flat of Kung Krabaen Bay, Thailand with notes on L. cf. corrugata. The Raffles Bulletin of Zoology. Suppl. 18: 57-73.
ROJWATTANA, P.. 2008: Population study of Laternula truncata (Bivalvia: Anomalodesmata: Laternulidae) in
the mangrove sand flat of Kung Krabaen Bay, Thailand with notes on L. cf. corrugata. The Raffles Bulletin of
Zoology. Suppl. 18: 57-73.
This plate presents two drifted pieces of wood discovered in the basal grind of the Kattendijk Sands Formation
(Lower Belgian Pliocene) at Doel (East Flanders, Belgium).
One of these piece of wood was completely disagreed, but one perfectly preserved tube of one teredinid bivalve
has resisted again the corrosion.
The scarcity of wood remains in all the Belgian Neogene strata prospected by the senior-author and his friends
could result from the poorness of the vegetal implantation along the coasts of this period.
Comments on Plate 33 and Plate 34
These plates presents photographs of some Belgian Pliocene shells of Corbula gibba (OLIVI, 1792), collected in
the upper part of the Merksem Sands Formation at Doel (Albert II dock), in the summer 2003.
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Their concentration rises up to twenty thousand individuals per square meter, and these ones were observable on
areas of more than fifty ares.
Dubblets and isolated valves of this species were concentrated in three successive sandy-glauconite levels, each
separated by ten centimetres of blue clay.
Each of these levels was separately prospected for its micro-teeth content. These sifting residues revealed the
presence of numerous Squalus acanthias, at least, four species of Raja sp. and thousands of rajiformes dermal
denticles.
The procentage of teeth of Raja sp. increased notably from the lower accumulation to the upper one.
This succession is the oldest Belgian wadden-sea-like deposit known.
These two plates presents two small septaria* reworked from the top of the Boom Clay Member. These were
discovered in the basal grind of the Edegem Sands Formation at Edegem (Antwerp Province, Belgium) during
the construction of the E3 Tunnel.
*These ‘septaria’ do not present ‘septae; they are in fact small and massive subspheric calcareous concretions concretions
which are fairly common, in the vicinity of Edegem, in the uppermost part of the Boom Clay.
Many of these septaria were bored by colonies of Martesia peroni COSSMANN & LAMBERT, 1844 which covered
the lower face of these concretions.
The whole surface of these septaria had been bored by numerous small Porifera of the Family Clionidae and by
some lithophagous Mollusca, such as these adult individuals of Martesia rugosa.
These plates regroup photographs of some individuals of Martesia rugosa BROCCHI, 1814 extracted from small
septaria disembedded, during the Lower Miocene, from the upper part of the Boom Clay Member.
After the dead of the individuals of this first population, the holes were inhabited by successive other populations
of Martesia rugosa, Saxicava arctica LINNAEUS, 1767 and Coraliophylla lithophagella (LAMARCK, 1819)*.
*According to the determinations of Dr. Arie Jansen (1964).
These three plates regroup diverse views and details of a large and unique fragment of the trunk of a palm-tree
discovered in the Imbrechts Sandpit at Neder-Okkerzeel (Flemish Brabant, Belgium), in the uppermost part of
the Brussels Sands Formation (Lutetian, Middle Eocene).
The tubes of the individuals of the very dense populations of teredinids present in this trunk fragment have some
morphological similarities with these of the extant Genus Uperotus GUETTARD, 1770.
Reference
TSUNODA, K. 1979: Ecological studies of shipworm attack on wood in the sea water log storage site.
Bulletin of the Wood Research Institute of the University of Kyoto. 65: 44 p.
This plate regroups some photographs of the larger enigmatica of the Invertebrate Collections of the I.R.S.N.B.,
which is presently abandoned on the public parking of this Institution.
For decencies, this strange and massive concretion was stored on the left side of the main entrance of the fossil
vertebrate’s room planified by Dr. Gilson and Dr. Dollo.
The senior-author has tried, without success, to discover from where could came this fossil labelled as gyrolithes.
Purchased by the Museum, gift of a private collector, donation of a Society or discovery, such a voluminous
fossil must legally have been registered, but no trace of this acquisition was mentioned in any registers of the
Museum*.
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*Till to 1948, the Royal Institute for Natural Sciences of Belgium has the status of a Museum.
Size, wegth, nature, walls, filling and epifauna
Size and weigth
This sandstone concretion has a height of 126 centimetres and 85 to 88 centimetres of diameter. Its weight was
estimated between 450 and 500 kilograms. I.R.S.N.B. was not in possession of an instrument allowing to precise
the weight of such an unstable mass when it was decided to displace this one.
Nature
It is not the shell, but the intern silicified mold of the habitat of a huge invertebrate, comprising three superposed
whole rooms and a last one presenting singular apertures.
Walls
The walls of the shelly habitat were, probably, dissolved by a chemical process which only allows to suppose
that their original thickness was comprised between 2,5 millimetres and 1 millimetre. Except for their sandstony
touch, their surfaces are smooth.
This thickness is suggested by the thin crystalline silicious substance which separates the successive rooms.
Filling
The fact the sediments filling the two lower rooms present an oblique stratification allows suggesting that, after
the dead of its builder, the shell was in an oblique position.
The filling of the shell is constituted by fine to very fine sands cores, interrupted by a very short phases of
income of coarser ones.
Epifauna
The highly damaged shelly remains of the invertebrate incrustations consist on carinated tubes of Polychaeta and
thecae of balanid Cirripedia. The tubes of carinated Polychaeta seem to be attributed to individuals of the Genus
Spirobrachus* de BLAINVILLE, 1818 (Family Serpulidae JOHNSTON, 1865) and the basal prints of diverse thecae
of balanid Cirripedia (Thoracia, Balaniformes) are of uncertain generic attribution.
*WoRMS 2015: Genus Pomatoceros PHILIPPI, 1844 is a synonym of Genus Spirobrachus de BLAINVILLE, 1818.
Other quite unperceptible and strongly abraded traces could be these of hydrozoan or bryozoan small colonies.
Some small perforations are also visible.
Today, all these invertebrates may be collected off, or along, the Atlantic * and the Mediterranean coasts, where
they live fixed on hard organic or inorganic supports at various depths.
*From northern Norway to southern Morocco.
All together, these data allow supposing that it was discovered in shallow marine waters
The presence of these extant taxa is not an argument allowing to attribute to this concretion an Holocene Age, it
only demonstrates that the extern side of this concretion was a perfect anchorage mass for diverse sessile inver
-tebrates during a part of this Period.
If it was dradged on the North Sea bottom, the most plausible zone is this where the streams of the Thames
River and the former Senne River intercrossed. The Hinder Banks* are a possible location point.
*See Max Poll 1947: Les Poissons marins de la Belgique: map 2.
Conclusion
None of these data allows proposing any age to this fossil.
Some possibile origins
A sedimentary level of a geological succession of a foreign country surrounding the North Sea, a concretion
trawled from the central part of the same Sea or a level located on another continent.
-99-
A concretion discovered in a foreign country surrounding the North Sea
Since the years 1850, all the fossiliferous strata of the countries surrounding the North Sea have been so
intensively prospected that it would be exceptional that similar fossils had never been observed or mentioned in
none of these.
A concretion trawled from the central part of the North Sea
The bottom of the central area of the southern part of the North Sea is constituted by an accumulation of massive erratic stone blocs produced by the smelting of the last icepacks which covered the Baltic countries.
This mass, called Eridan, contains diverse types of boulders of which some have a sedimentary origin. This
concretion could have been extracted of one of these sedimentary levels.
But, no similar concretion had been discovered in none of the sedimentary levels surrounding the Baltic Sea, an
area so intensively prospected.
A concretion carried by ship from another continent
Another possibility is that the size and the morphology of this concretion have surprised a naturalist during his
exploration of a far country and that he decided to ship this one to a European Institution.
Violent storms are common events in the North Sea * and the number of sinked ships by these storms is very
high. It is possible that the ship carrying this enigmatic concretion sunk during one storm and those fishermen
discovered it in their net. Conscients of the singularity of their discovery, they give it to a man they appreciated:
Dr. Gilson.
*See: BAYENS, E. 2002: De verraderlijke Zee - Scheepsrampen in de Noordzee. 256 p. Lannoo. ISBN: 90 209 4997 4
A falsification
False fossils have sometimes been produced as a joke* to test the knowledge of some pretentious specialists, but
such a singular mass had none interests for any world-fame Belgian paleontologists of this period.
*Such as the man of Pildown.
Conclusion
The geographical origin and the nature of this huge concretion remain mysteries.
It becomes urgent to protect this unique ichnofossils again an ineluctable detoriation.
-100-
16. Acknowledgements
The authors are particularly greatfull to
Anne ABREAR (AU), Lucia BORGES (D), Pieter DE SCHUTTER (B),
Hugues DOUTRELEPONT (B), Robert MARQUET (B), Catarina PIMIENTO (U.S.A.),
Guido and Philippe POPPE (B), Carsten RENKER (D), Janet VOIGTH (U.S.A.),
Olev VINN (EE) and Argyros ZENETOS (EL).
for their precious help.
-101-
The original PDF was sent, the 2 August 2015
to the Belgian royal Library,
Legal Electronic dépôt Survey
and, because of techinal problems, the 3 August 2015
to the following Institutions:
In Belgium:
I.R.S.N.B., S.G.B., U.L.B., V.U.B., R.U.G., U.Lg.,
M.R.A.C., K.M.C.A.,
Musée de Zoologie de l’U.L.B. and Musée d'Histoire naturelle de Mons.
In other Countries:
N.M.N.H. (GB), B.G.S. (GB), B.M.N.H. (GB),
Naturalis Biodiversity Center Leiden (NL),
M.N.H.N. (F), Musée d’Histoire Naturelle de Lille (F),
Krahuletz-Museum Eggenburg (A), Bergbaumuseum Klagenfurt (A),
Biozentrum Grindel und Zoologisches Museum, Hamburg (D),
Geologisches Museum München (D), Niedersächsisches Landesmuseum Hannover (D),
Museum für Naturkunde, Berlin (D), Senckenberg Museum, Frankfurt (D),
A.M.N.H. (USA), Auckland Museum (NZ), Field Museum of Chicago(USA),,
KARLSRUHE Museum (D), Alexander von Humboldt Museum (D),
and Museum de Genève (CH).
And to the following colleagues and former field-collaborators:
Alroy John, Amaru Kazuto, Baut Jean-Paul,
Barrull John, Boel Jacques, Bonde Niels, Bor Taco,
Boulvain Frédéric, Bourdon Jim, Bosselaers Mark, Case Gerard-Ramon,
Collier Eric, Cunnny Gilles, Doutrelepont Hugues, Dusar Michiel, Dutheil Didier, Finet Yves,
Fraije René, Garot Philippe,Gonzalez Gerardo, Grogan Ellen,
Guenegues Serge, Hooker Jerry, Hovestadt Dirk, Kitamura Naushi, Last Peter,
Lenglet Georges, Louwye Stephen, Lowe Ruth, Kedra Marika, Kriwet Jürgen, Kukuev Efy,
Kupryanova Elena, Ishihara Hajima, Malakovski Svetlana, Migom Frédéric, Mollen Frederik,
Pfeil Fritz, Pöllerspock Jürgen, Popov Evgeny, Poulicek Mathieu, Préat Alain,
Rac-Lorenz Helen, Rees Tony, Romero Javier, Sabourin Nadine, Sebastiani Didier, Steurbaut Etienne,
Sigurdsson Stein, Stehmann Matthias, Schlögel Jan, Telesh Irina,Tabachnikov Konstantin,
Takada Koji, Tarifa Silva Eduardo, Taverne Louis, Taylor Paul, Thies Detlev,
Udovichenko Nikolaï,Underwood Charlie, Van Bakel, Barry, Vandenberghe Noël,
van Goethem Jackie, Viktoras Didziulis,Vinn Olev,
Yabumoto Yoshikata,Ward David, Welton Bruce, Wesselingh Frank,
Wille Eric,Winderickx Didier, Wirtz Peter and Zuber Pavel.
Editeur responsable: Docteur Jacques Herman. I.S.S.N. : 2033 - 6365
Beigemsesteenweg 319. 1852 Beigem (Grimbergen).
Belgique - België - Belgien.
G-mail: jacquesalbertherman@gcom.be
Website, freely accessible: www.geominpal.be
-102-

Addition 2 to Géominpal Belgica. 5.2.: Presence of tubes of teredinid

Transcription

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