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Verlag Dr. Friedrich Pfeil · München

Verlag Dr. Friedrich Pfeil

Guide to the Identification of

MarineMeiofauna

edited by

Andreas Schmidt-Rhaesa

Guide to the Identification of

MarineMeiofauna

edited by

Andreas Schmidt-Rhaesa

Marine meiofauna, the community of smallest animals in marine sediments, is a fascinating and important part of the marine ecosystem. In this book, 53 authors introduce the 32 animal groups that occur in the meiofauna. Chap-ters contain information on where and how to sample meiofaunal animals as well as how to identify them. Keys help in the identification.

This book complements existing literature on meiofauna and may serve as a fascinating introduction into meio-faunal taxa as well as a practical guide to work with meio-fauna in the field or in the laboratory.

The rich illustration of the book and the expertise of spe-cialists make this book suitable for beginners as well as experienced researchers on meiofauna.

Verlag D

r. Friedrich Pfeil

Guid

e to the Id

entification o

f

Marine

Meio

faunaed

ited b

y

And

reas Schm

idt-R

haesa

92

A

50 µm

B

pr

teph

mco

C

eyey

ey

ey

pr

te

ph

mco

50 µm

100 µm

Fig. 6.42. Rhabdocoela, Kalyptorhynchia, Eukalyptorhynchia, Nannorhynchididae. A. Toia sp. from Waquoit Bay, MA, USA, showing proboscis (pr), pharynx (ph) and position of male organ (mco). B. Anterior end of above specimen, showing pigmented eyes (ey) with lenses. Both images courtesy of Dr. Christopher Laumer. C. Nan-norhynchides sp. from British Columbia, Canada, showing the proboscis, pharynx, paired testes (te), eyes with lenses, and the male copulatory organ.

Dr. Marco Curini-Galletti is warmly thanked for editing the Proseriata section of the key.

References

Appeltans, W. et al. (2012): The magnitude of global marine species diversity. Current Biology 22: 2189-2202.

Guide to the Identification of Marine Meiofauna

108

Only in Xenodasys, epidermal cells fill the interior of the cuticular structures (Rieger & Rieger 1977). There are two types of glandular epidermal structures: (a) paired, single-celled epidermal glands (e. g., Figs. 7.5G-H, 7.7F) of unknown func-

tion in rows along the dorsal body wall (Kieneke & Schmidt-Rhaesa 2015) and (b) adhesive tubes, either in different positions and numbers along the body of macrodasyid gastrotrichs or only as a single posterior pair in most Paucitubulatina

head-neck-

trunk region

furca

furca

pharyngeal regionintestinal region

head

-ne

ck-

trun

k re

gion

pedicles

phar

ynge

al r

egio

nin

test

inal

reg

ion

A B

CTbA

TbVLoo tes

tesoo

ovxo

cir

sci

sci

cgdco

ph

pp

PhJIn

fo

co

eg

ag

ag

pn

TbP

TbP

ag

ph

phso

ahso

ov

cgdco

vd

mig

TbVL

mig

vncvnc

Fig. 7.4. Schematic body organization and morphology of marine Gastrotricha. A. Generalized macrodasyid gastrotrich, reproductive and nervous systems. B. Protonephridia, intestinal system, epidermal and adhesive glands. C. Generalized xenotrichulid gastrotrich. Abbreviations: ag, adhesive glands; ahso, anterior head sen-sory organ; cg, cerebral ganglion; cir, sensory cirri; co, caudal organ; dco, dorsal commissure; eg, epidermal glands; fo, frontal organ; mig, midgut; oo, mature oocyte; ov, ovaries; ph, pharynx; PhJIn, pharyngeo-intesti-nal junction; phso, posterior head sensory organ; pn, protonephridia; pp, pharyngeal pores; sci sensory cilia; TbA/TbP/TbVL, anterior/posterior/ventrolateral adhesive tubes; tes, testes; vd, vas deferens; vnc, ventral longitudinal neurite cords; xo, x-organ. A & B dorsal views, redrawn from Ruppert 1991, C dorsal view, redrawn from Kieneke & Schmidt-Rhaesa 2015.

Guide to the Identification of Marine Meiofauna

419

A

FE

D

B

50 µ

m

20 µm

20 µ

m

20 µ

m

50 µ

m

mumuto

ps

ps to

as

mc

add

mc

mt

or

cs

vpvlp

ip

10 µ

m

flN

ts

C

lp

22 Loricifera

247

100 µm

A

mg

B

FE

C

D

100 µm

50 µm

ey

mo

mg

gl

ba

pa ba

ch

ch

ch

ch

ba

100 µm

100 µm

zo

200 µm

100 µmG H 100 µm

10 µm

ph

mg

mgmg

br

dc

cr

dc

Fig. 11.3. Fauveliopsidae (A), Cirratulidae (Ctenodrilinae) (B-C, F), Acrocirridae (D), Psammodrilidae (G-H). A. Fauveliopsis sp. (photo: A. Martinez). B. Ctenodrilus sp. (photo: G. Rouse). C. Ctenodrilus sp. D. Macrochaeta sp. E. Macrochaeta sp. F. Raphidrilus sp. G. Psammodrilus moebjergi Worsaae & Sterrer, 2006. H. Psammodrilus norenburgi Worsaae & Martínez, 2018 (insert: close-up of hooked chaetae (uncini)). Abbreviations: ba, branchia; br, brain; ch, chaeta; cr, collar region; dc, dorsal cirrus; ey, eye; gl, gland; mg, midgut; mo, mouth; pa, palp; ph, pharynx; zo, zooid.

T. ellenscrippsae Kerbl, Vereide, Gonzalez, Rouse & Worsaae, 2018; T. windansea Kerbl, Vereide, Gonzalez, Rouse & Worsaae, 2018.

References. Harmer (1889), Jägersten (1944), Westheide (1967, 1985, 2008, 2016), Riser (1999), Simonini et al. (2003), Kajihara et al. (2015), Kerbl et al. (2016, 2018).

11 Annelida

Guide to the Identification of

MarineMeiofauna

edited by

Andreas Schmidt-Rhaesa

608 pages198 coloured and 135 black-and-white figures,

6 tables, 40 partly illustrated identification keys

24.5 × 17.3 cmHardcover

ISBN 978-3-89937-244-168.00 ˜

Contents

Foreword (pp. 7-8)

1 Introduction (pp. 9-16)2 General key (pp. 17-23)3 Ciliophora Stephen C. Landers (pp. 24-32)4 Cnidaria Andreas Schmidt-Rhaesa, Sofia Pyataeva and

Allen G. Collins (pp. 33-43)

5 Acoelomorpha Ulf Jondelius and Ylva Jondelius (pp. 44-53)6 Platyhelminthes Julian P. S. Smith III, Niels Van Steenkiste

and Tom Artois (pp. 54-103)7 Gastrotricha Alexander Kieneke, Lukas Münter and Ole

Riemann (pp. 104-163)8 Rotifera Wilko Ahlrichs and Ole Riemann (pp. 164-198)9 Gnathostomulida Martin V. Sørensen and Wolfgang Sterrer

(pp. 199-226)10 Nemertea Jörn von Döhren and Thomas Bartolomaeus

(pp. 227-238)11 Annelida (excluding Clitellata and Sipuncula) Katrine

Worsaae (pp. 239-270)12 Oligochaeta (Clitellata) Olav Giere (pp. 271-276)13 Sipuncula Anja Schulze, Gisele Y. Kawauchi and Alvaro

Migotto (pp. 277-284)14 Entoprocta Claus Nielsen (pp. 285-287)15/16 Mollusca (Gastropoda and Aplacophora) Katharina M.

Jörger (p. 288)15 Gastropoda Katharina M. Jörger, Timea P. Neusser, Bastian

Brenzinger and Michael Schrödl (pp. 289-307)16 Aplacophoran molluscs: Solenogastres and Caudofoveata

Franziska S. Bergmeier and Katharina M. Jörger (pp. 308-320)17 Bryozoa Jörg Ott and Thomas Schwaha (pp. 321-326)18 Brachiopoda Carsten Lüter (pp. 327-330)19 Nematoda Nicole Smol, Tânia Nara Bezerra and Wilfrida

Decraemer (pp. 331-386)20 Priapulida Andreas Schmidt-Rhaesa (pp. 387-390)21 Kinorhyncha Martin V. Sørensen and Fernando Pardos

(pp. 391-414)22 Loricifera Shinta Fujimoto and Reinhardt Møbjerg Kristensen

(pp. 415-427)

23 Tardigrada Jesper Guldberg Hansen and Reinhardt Møb-jerg Kristensen (pp. 428-444)

24 Halacaridae (Prostigmata: Acari) Ilse Bartsch (pp. 445-453)25 Mystacocarida Jørgen Olesen (pp. 454-458)26 Cephalocarida Jørgen Olesen (pp. 459-464)27 Copepoda Kai Horst George, Sahar Khodami, Terue

Cristina Kihara, Pedro Martínez Arbizu, Alejandro Martínez, Nancy F. Mercado-Salas, Karin Pointner and Gritta Veit-Köhler (pp. 465-533)

28 Ostracoda Dietmar Keyser (pp. 534-560)29 Isopoda Nicole Coineau (pp. 561-576)30 Cumacea Ute Mühlenhardt-Siegel (pp. 577-581)31 Chaetognatha Andreas Schmidt-Rhaesa (pp. 582-584)32 Enteropneusta Sabrina Kaul-Strehlow and Katrine Wor-

saae (pp. 585-587)33 Echinodermata: Holothuroidea David L. Pawson (pp. 588-

591)34 Tunicata (Ascidia) Andreas Schmidt-Rhaesa (pp. 592-597)

My meiofauna story How (not) to deal with sponsors (Andreas Schmidt-Rhaesa)

(p. 598) How I came to be a meiobenthologist (Jörg Ott) (p. 599) Fantastic Meiofauna or Meiofauna at the Verge of Cred-

ibility (Olav Giere) (pp. 599-600) Sampling Symsagittifera: Green colour disappearing into

the sand (Andreas Schmidt-Rhaesa) (pp. 600-601) Serendipity – or how I stumbled onto gnathostomulids

(Wolfgang Sterrer) (pp. 601-603) The “archiannelid” lump (Carsten Lüter) (pp. 603-604) A sieve for megafauna (Andreas Schmidt-Rhaesa) (p. 604)

Authors (pp. 605)

294

Fig. 15.2. Overview of the different lineages of meiofaunal slugs: Rhodopemorpha (A-B), Nudibranchia (C-D) and Cephalaspidea (E-G). A. Rhodope sp. from Hawaii. B. Helminthope sp. (‘Crucispiculata-form’) from Hawaii. C. Embletonia pulchra from Mediterranean. D. Pseudovermis salamandrops from Brazil. E. Philine sp. from Hawaii. F. Philinoglossa sp. from Indonesia. G. Pluscula cuica from Panama.

A

B

C D

E

F G

500 µm

A

B

C D

E

F G

500 µm

Guide to the Identification of Marine Meiofauna

354

A B

am

C D

Fig. 19.19. Triplonchida: Triodontolaimidae: Triodontolaimus: A. View on one tooth with denticles. B. Amphid (am). Pandolaimidae: C. Buccal cavity and separate rows of outer labial papillae and cephalic sensillae in Pan-dolaimus. Rhabdodemaniidae: D. View on teeth in Rhabdodemania. A, B: ©Alessandra Botelho.

Guide to the Identification of Marine Meiofauna

462

A

B

C

D

A

B

D

C

Antenna 2, exopod

Antenna 1

50 µm

50 µm100 µm

ts7

ts1

ts9

ts15

ts19

Telson

Thoracopods

Egg carrier limb

ts8 - thoracopod absent

Cephalic shield

Mandible

Maxillae 1

Hypostome/clypeus

Labrum

Antenna 1

Antenna 2

Thoracopod 7

ts8 -

thoracopod absent

Egg carrier limb

Maxilla 2

Thoracopod 1

Thoracopod

“Pseudepipod”

Exopod

Endopod

THORAX

Furcal rami

Atrium oris

Antenna 2

Antenna 2, endopod

Endites

Tergopleurae100 µm

ABDOMEN

Antenna 2, exopod

Antenna 1

50 µm

50 µm100 µm

ts7

ts1

ts9

ts15

ts19

Telson

Thoracopods

Egg carrier limb

ts8 - thoracopod absent

Cephalic shield

Mandible

Maxillae 1

Hypostome/clypeus

Labrum

Antenna 1

Antenna 2

Thoracopod 7

ts8 -

thoracopod absent

Egg carrier limb

Maxilla 2

Thoracopod 1

Thoracopod

“Pseudepipod”

Exopod

Endopod

THORAX

Furcal rami

Atrium oris

Antenna 2

Antenna 2, endopod

Endites

Tergopleurae100 µm

ABDOMEN

Fig. 26.2. Morphology of Lightiella monniotae. A. Lateral view of entire adult specimen. B. Ventral view of labrum and cephalic appendages. Maxillae 2 have been removed. C. Thoracopod seen from posterior. D. Ventral view of cephalon and thorax showing labrum and cephalic and thoracic appendages.

Guide to the Identification of Marine Meiofauna

409

C DA B

G H

E F

I J

21 Kinorhyncha

583

A

B

C D

at

gs

gs

at

gs

Fig. 31.2. Spadella kappae. A. Head showing spread grasping spines (gs) and the anterior teeth (at). B. Scanning electron micrograph of the head from frontal, showing the same structures as labelled in A. C. Eyes. D. Unspread grasping spines close to the head.

2005, S. kappi Schmidt-Rhaesa & Vieler, 2018 and S. nunezi Casanova & Moreau, 2004.

Geographic distribution

The five species considered here have a very patchy distribution and were found on single oc-casions. Spadella boucheri is from Japan, S. gaetanoi is from Hawaii, S. nunezi was collected on the Canary Islands (Lanzarote), S. interstitialis is from Italy (Elba and Pianosa) and S. kappi is from France (Roscoff).

Conditions where the taxon is found

The five species were collected from sandy sub-strates and seem to prefer coarse sand (such as “Amphioxus”-sand). Some species were collected in the beach region, others from sediments down to 70 m.

Preferred extraction method

Specimens were collected by different methods, such as direct sampling from formalin-fixed sediment samples (with or without rose-bengal staining), by Ludox extraction from formalin-fixed samples or by narcotization with MgCl2.

31 Chaetognatha

589

33 Echinodermata: Holothuroidea

Fig. 33.1. A. Rhabdomolgus ruber Keferstein 1862 (after Becher, 1907). B. ? Patinapta ooplax (Marenzeller) (after Rao 1973). C. Anchor and anchor plate from Labidoplax buski McIntosh (after Salvini-Plawen 1972b). D. Wheel from Trochodota furcipraedita Salvini-Plawen, 1972 (after Salvini-Plawen 1972a). E. Auricularia nudibranchiata Chun (after Pawson 1971). F. Wheels from Myriotrochus vitreus (Sars) (after Salvini-Plawen 1977). G. Synaptid (?) juve-nile from Bahamas (provided by A. Schmidt-Rhaesa).

A B

C

D

E

F

G

30 µm

30 µm

30 µm30 µm